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MUTATIONS,  VARIATIONS,  AND  RELATION- 
SHIPS OF  THE  OENOTHERAS 


BY 


D.  T.  MACDOUGAL,  A.  M.  VAIL.  AND  G.  H.  SHULL 


WASHINGTON,  D.  C: 
Published  by  the  Carnegie  Institution  of  Washington 

1907 


North  Carolina  State  Library 


Gift  of 


North  CaroliDMRate  ua^aty 


MUTATIONS,  VARIATIONS,  AND  RELATION- 
SHIPS OF  THE  OENOTHERAS 


BY 


D.  T.  MACDOUGAL,  A.  M.  VAIL,  AND  G.  H.  SHULL 


WASHINGTON,  D.  C. : 

Published  by  tlie  Carnegie  Institution  ot  Washington 

1907 


/ 

/ 


Carnegie  Institution  of  Washington  Publication  No.  8i. 
Papers  of  the  Station  for  Experimental  Evolution,  No.  q. 


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MUTATIONS,  VARIATIONS.  AND    RliLATIONSHlPS 

OP  THE  OHNOTHHRAS. 


By  D.  T.  MacDougau,  A.  M.  Vail,  and  G.  H.  Shull. 


SCOPE  OF  INVESTIGATION. 

The  Oenotheras  have  furnished  so  much  evidence  of  importance  in  connec- 
tion with  saltatory  action  in  heredity  that  it  has  been  deemed  important  to 
continue  the  cultural  investigation  of  the  group  begun  in  1902. 

Sseds  representing  the  spscies  in  cultivation  in  the  principal  botanical 
gardens  of  the  world  have  been  procured,  and  these,  in  addition  to  a  large  num- 
ber of  forms  native  to  eastern  North  America,  have  been  grown  in  guarded 
cultures. 

Attention  has  bsen  paid  to  the  occurrence  of  mutants  in  Oenothera  lamarck- 
ian2  with  a  view  to  testing:  the  coefficient  of  mutabilitv  and  the  influence  of 
environmental  conditions  on  mutation.  Extensive  sowings  have  been  made 
for  the  purpose  of  finding  derivatives  hitherto  undetected,  with  a  coefficient  of 
mutability  so  small  as  to  have  escaped  observation.  Descriptions  of  known 
mutants  have  been  made  independently  for  the  purpose  of  comparison  with 
supposedly  identical  forms  in  Amsterdam  and  of  facilitating  observations 
of  all  kinds  upon  the  subject. 

Many  important  relations  between  mutants  and  their  parents  may  be  most 
advantageously  considered  by  statistical  methods,  and  the  studies  begun  by 
one  of  the  authors  in  1904  have  been  continued  and  extended  to  include  addi- 
tional mutants.  The  height  and  branching  of  the  stems  and  the  width  and 
length  of  the  leaves  have  been  again  taken  into  account,  but  owing  to  the 
great  susceptibility  of  these  organs  to  variation  in  direct  response  to  environ- 
ment, measurements  have  also  been  made  upon  the  buds.  The  lesser  varia- 
tion of  the  latter  in  correlation  with  vegetative  characters  makes  them  much 
more  satisfactory  for  the  study  of  hereditary  relations,  and  it  is  clear  that 
their  statistical  study  in  connection  with  pedigree-cultures  will  demonstrate 
in  several  generations  the  permanence  or  evanescence  of  the  mutant  types 
and  give  decisive  answers  to  such  questions  as  the  relation  between  fluctua- 
tion and  mutation  and  the  "fixing"  of  variations  through  self-fertihzations 
or  their  disappearance  through  crossings. 


2        MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 

Guarded  psdigree-cultures  have  been  made  from  pure  seeds  of  native  species 
of  evening- primroses  which  had  not  previously  been  brought  under  close 
observation,  with  a  view  to  procuring  additional  evidence  on  mutabiHty  in 
this  group. 

The  preliminary  examination  made  it  apparent  that  the  group  comprises  a 
swarm  of  clearlv  separable  species,  many  of  which  had  not  been  recognized 
by  the  taxonomists  and  which  gave  basis  for  the  current  opinion  as  to  the 
wide  variability  of  the  common  evening-primrose  (D.  biennis). 

The  studv  of  genetic  relationships  and  of  the  phenomena  of  hybridization 
in  general  has  necessitated  the  organization  of  extensive  cultures,  which  wall 
need  continuance  for  two  or  more  seasons  before  definite  results  may  be 
announced,  and  their  discussion  is  reserved  for  a  future  paper. 

A  few  cases  of  fixed  hybrids  have  been  encountered,  however,  in  which  the 
crossing  of  two  species  resulted  in  the  production  of  a  progeny  consisting  of 
one  fixed  tvpe  constant  in  successive  generations.  Forms  arising  in  this  way 
are,  in  reality,  new  species,  and  doubtless  many  such  have  arisen  naturally 
and  now  constitute  a  part  of  the  native  flora. 

Striking  cases  of  vegetative  mutation,  or  bud-sports,  having  arisen,  atten- 
tion has  been  devoted  to  a  study  of  the  inheritance  of  the  saltatory  groups  of 
characters. 

A  systematic  attempt  to  localize  mutatory  changes  in  the  life-cycle  of  the 
sporoph}^e  has  been  made,  and  has  met  with  marked  success  in  a  preliminary 
wav,  since  it  has  been  possible  to  induce  new  mutants  by  the  use  of  solutions 
of  strong  osmotic  activity  and  by  highly  dilute  preparations  of  mineral  salts, 
some  of  which  are  poisonous  to  plants  in  high  concentrations  and  stimulative 
in  low  concentrations.  Finally,  an  attempt  has  been  made  to  bring  the  facts 
disclosed  in  the  descriptive  part  of  the  paper  into  correlation  with  prevailing 
theories  as  to  phylogeny  and  evolutionary  procedure. 

The  principal  results  were  obtained  from  cultural  investigations  in  the 
experimental  grounds  and  greenhouses  of  the  New  York  Botanical  Garden 
and  of  the  Station  for  Experimental  Evolution  at  Cold  Spring  Harbor,  Long 
Island,  and  were  also  extended  in  the  later  stages  to  the  Desert  Botanical 
Laboratory  of  the  Carnegie  Institution  of  Washington  at  Tucson,  Arizona, 
thus  securing  the  advantages  of  a  wide  range  of  climate  and  soils. 

A  note  regarding  the  various  mutants  was  presented  at  the  weekly  botanical 
convention  at  the  New  York  Botanical  Garden,  November  i,  1905,  and  a 
second  one,  dealing  especially  with  sports  or  vegetative  mutations,  before  the 
Torrey  Botanical  Club,  on  November  4,  1905.  A  lecture  dealing  with  his- 
torical aspects  of  the  phases  of  the  subject  under  investigation  was  presented 
to  the  Barnard  Botanical  Club,  December  18,  1905,  in  which  the  first  announce- 
ment was  made  of  the  induction  of  mutants  by  chemically  and  osmotically 
active  stimuli.     A  full  description  of  the  technique  of  pedigree-cultures  and 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS.        3 

of  the  methods  emploved  in  the  stimulation  of  ovaries  was  given  in  a  lecture 
at  the  Woods  Hole  Marine  Biological  Laboratory,  July  20,  1906,  in  conjunction 
with  which  derivatives  thus  obtained  of  the  second  generation  in  Raimannia 
and  of  the  first  of  0.  biennis  were  exhibited.  The  results  of  Dr.  Shull's 
statistical  inquiries  were  presented  before  the  Botanical  Society  of  America  at 
New  Orleans,  January  3,  1906,  and  various  notes  have  been  presented  before 
other  societies. 

PEDIGREE-CULTURES  OF  OENOTHERA  LAMARCKIANA. 

Arrangements  were  made  by  which  seeds  were  obtained  from  four  separate 
individuals,  purely  fertilized  with  their  own  pollen.  This  was  done  by  apply- 
ing the  pollen  to  the  pistil  of  the  same  flower  or  of  another  flower  of  the  same 
plant.  All  sowings  were  made  in  earthenware  pans,  30  by  30  cm.,  filled  with 
soil  sterilized  in  an  autoclave  as  in  previous  cultures,  and  when  sufficiently 
developed  the  seedlings  were  removed,  to  be  transplanted  or  discarded,  as 
the  conditions  of  the  experiments  demanded.  The  cultures  are  given  under 
the  key-numbers  by  which  they  were  designated  in  the  journal  of  the  experi- 
mental garden. 

A.  I.  0. — -A  packet  of  seeds  which  was  given  the  foregoing  designation, 
which  had  bsen  harvested  in  Amsterdam  in  1901,  and  was  obtained  directly 
from  Professor  Ds  Vries.  The  sowing  was  made  in  August,  1904,  and  the 
small  rosettes  were  inspected  by  Professor  De  Vries  late  in  September,  1904, 
and  he  kindly  assisted  in  the  identification  of  a  few  of  the  mutants  included. 
Some  confusion  in  the  record  makes  it  impossible  to  give  the  exact  census  of 
the  culture,  but  it  comprised  between  500  and  600  seedlings,  among  which 
26  mutant  derivatives  were  identifiable,  and,  so  far  as  possible,  two  represent- 
atives of  each  type  were  transplanted  to  the  experimental  garden  in  May, 
1905,  coming  into  bloom  about  60  days  later. 

The  authors  have  not  had  the  opportunity  of  inspecting  living  specimens 
of  all  of  the  mutants  which  have  appeared  in  Amsterdam  and  which  have 
been  described  by  Professor  De  Vries,  and  of  the  9  forms  seen  but  5  have 
been  conclusively  recognized.  Of  these,  O.  oblonga  was  represented  by  12 
individuals,  constituting  46  per  cent  of  the  total  number  of  mutants.  The 
average  frequency  of  this  type  among  the  mutative  progeny  is  i  per  cent,  or 
20  per  cent  of  the  mutants,  although  in  one  series  De  Vries  found  176  ohlongas 
among  a  total  of  334  mutants  derived  from  a  culture  consisting  of  over  14,000 
individuals.  In  this  instance  oblonga  constituted  nearly  53  per  cent  of  the 
mutants  or  1.25  per  cent  of  the  entire  progeny.  (In  De  Vries,  1905,  p.  545, 
the  proportion  of  oblonga  in  cultures  is  given  as  10  per  cent,  which  is  a  mis- 
print for  I  per  cent.) 

In  the  New  York  culture  under  discussion,  lata  was  represented  by  one 
individual,  nanella  by  one,  albida  by  two,  and  gigas  by  one.     Of  these,  the 


4        MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE)    OENOTHERAS. 

appearance  of  gigas  is  the  most  notable,  since  it  has  occurred  but  few  times 
in  the  last  twenty  years  in  pedigree-cultures  of  0.  lamarckiana. 

The  remaining  four  forms  among  the  mutants  were  represented  by  one, 
one,  two,  and  five  individuals.  Of  these,  one  type  with  orbicular  leaves 
presented  a  most  striking  departure  from  the  parental  form,  but  failed  to 
perfect  flowers,  so  that  but  little  information  concerning  it  was  gained.  The 
form  in  question  was  reprseented  by  one  individual  only,  and  did  not  appear 
in  anv  other  culture 

It  is  to  be  noted  that  the  seeds  used  in  this  culture  had  been  stored  for  three 
years.  Doubtless  the  parental  type  and  the  derivatives  are  characterized 
by  varying  and  different  powers  of  endurance,  so  that  the  coefficient  of  fre- 
quency of  the  several  forms  in  question  might  be  expected  to  be  slightly 
dift'erent  from  that  found  in  freshly  harvested  seeds. 

C.I.  2. — A  lot  of  purely  fertilized  seeds  derived  from  one  individual,  har- 
vested in  the  New  York  Botanical  Garden  in  1903,  was  sown  in  sterilized  soil 
in  the  greenhouse  in  August,  1904.  In  September  the  plantlets  w^ere  so  far 
advanced  as  to  admit  of  recognition  of  a  number  of  forms,  and  a  number  of 
duplicates  of  the  parental  type  were  discarded.  This  process  was  repeated 
at  intervals  throughout  the  winter,  and  the  remaining  rosettes  were  trans- 
planted to  the  experimental  garden  in  May,  1905.  vSeven  derivatives  were 
seen,  including  33  individuals.  Among  these,  were  2  of  scintillans,  3  of  albida, 
18  of  oblonga,  and  2,  2,  3,  and  3  of  four  other  unknown  types.  Some  of  these 
unknown  types  also  occurred  in  other  cultures,  and  3  of  the  known  and 
unknown  were  not  duplicated  in  other  progenies. 

The  total  number  of  the  seedlings  included  was  probablv  not  much  above 
500,  and  if  this  were  conclusively  confirmed  it  would  furnish  an  illustration  of 
a  progeny  in  which  the  mutants  constituted  about  6  per  cent  of  the  entire 
number.  Unfortunately  some  confusion  was  found  in  the  record.  Before 
this  was  discovered,  the  senior  author  announced  that  he  had  succeeded  in 
modifying  the  coefficient  of  mutability,  but,  as  is  to  be  seen  from  the  above, 
the  evidence  is  not  conclusive.  The  large  proportion  of  oblonga  is  noticeable, 
and  this  form  constitutes  54  per  cent  of  the  total  number  of  the  mutants. 

D.  I.  7. — A  lot  of  purely  fertilized  seeds  taken  from  one  individual,  grown 
in  the  New  York  Botanical  Garden  in  1904,  was  sown  in  the  garden  in  August, 
1904,  furnishing  a  total  of  499  plantlets.  Development  proceeded  so  rapidly 
that  it  was  possible  to  select  and  discard  244  duplicates  of  the  parental  type 
on  December  4,  1904.  The  remaining  ones  were  transferred  to  small  pots. 
Forty-three  more  duplicates  were  discarded  on  Februarv  9,  1Q05,  34  on  Feb- 
ruary 18,  40  on  March  3,  6  on  INIarch  21,  2  on  March  24,  3  on  March  27,  7  on 
March  30,  and  i  on  April  17.  On  May  15,  19  mutants  and  2  of  the  parental 
type  were  transplanted  to  the  experimental  garden.  The  detail  given  above 
illustrates  the  general  procedure  in  all  such  cultures. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.        5 

As  a  result  of  the  continued  inspection  of  the  cuhure  it  was  seen  to  include 
3  of  nanella,  2  of  lata,  6  of  albida,  and  3  of  oblonga,  while  the  remainder  could 
not  be  identified.  One  was  purely  pollinated  to  test  its  relationship  with 
gigas.  The  coefficient  of  mutability  in  this  culture  was  seen  to  be  barely  3 
per  cent,  and  the  proportion  of  oblonga  was  much  lower  than  in  any  other 
culture. 

D.  I.  9. — A  lot  of  purely  fertilized  seeds  designated  as  above,  harvested  in 
the  New  York  Botanical  Garden  in  August,  1904,  was  sown  in  two  earthen- 
ware pans  filled  with  sterilized  soil,  early  in  November,  1904;  604  plantlets 
were  produced ;  390  duplicates  of  the  parental  type  were  discarded  on  Decem- 
ber 4th,  1904,  and  66  on  December  15th;  77  were  transplanted  for  further 
inspection  on  December  21st,  and  the  remaining  71  on  January  15th,  1905;  of 
these,  59  were  discarded  on  February  9th,  33  on  February  i8th,  15  on  March  3d, 
3  on  March  21st,  5  on  jMarch  24th,  and  4  on  March  30th;  19  mutant  individuals 
were  transplanted  to  the  experimental  garden  on  May  i6th,  1905,  of  which 
I  was  nanella,  2  were  lata,  2  scintillans,  i  albida,  and  2  oblonga,  and  the 
remainder  were  of  types  not  identified  by  the  authors. 

The  coefficient  of  mutability  in  this  instance  is  seen  to  be  about  3  per  cent, 
and  that  of  oblonga  very  small  in  comparison  with  the  general  frequency  of 
this  derivative. 

IDENTITY  AND  DISTRIBUTION  OF  OENOTHERA  LAMARCKIANA. 

Since  the  cultures  of  the  evening-primroses  was  begun,  a  few  years  since, 
no  opportunity  has  been  neglected  to  attempt  to  trace  0.  lamarckiana  to  its 
original  habitat  and  to  establish  its  relationship  to  other  species  of  the  genus. 
Among  numerous  bibliographical  discussions  of  interest  in  this  connection, 
one  by  Miller  (1760)  is  of  great  interest.  He  says  regarding  the  "Tree  Prim- 
rose with  oval  spear-shaped  indented  Leaves,  and  Flowers  proceeding  from 
the  wings  of  the  Teaves  on  the  Upper  Part  of  the  Stalk:" 

This  plant  is  also  a  Native  of  North  America;  but  was  the  first  species  of  the  Genus  which 
was  brought  to  Europe,  so  it  is  more  commonly  seen  in  the  Gardens  than  any  of  the  other 
species.  In  some  parts  of  Europe,  this  is  spread  about  from  the  Gardens  in  such  plenty, 
that  it  might  be  supposed  a  native  there.  In  a  small  wood  near  Haarlem  in  Holland,  this 
plant  covered  the  ground  insomuch  that  n^any  skilful  persons  supposed  it  was  a  native  of 
that  place.  But  it  may  be  easily  accounted  for,  because  the  gardeners  who  live  near  that 
place  are  chiefly  .^orists,  and  they  annually  change  the  earth  of  the  beds  in  their  gardens; 
so  by  carrying  out  of  their  old  earth  from  their  beds,  in  which  many  of  the  seeds  were 
scattered,  the  plants  came  up  there;  and  those  being  suffered  to  scatter  their  seeds,  had 
filled  the  whole  wood  with  the  plants. 

This  differs  from  tlie  first  sort  (described  and  figured  as  0.  biennis)  in  having  broader 
leaves;  the  stalk  grows  taller,  and  the  flowers  are  much  larger.  Both  these  sorts  will  thrive 
in  the  Smoak  of  London  better  than  most  plants. 

The  appended  descriptions  and  the  plate  (No.  189,  dated  1757)  very  fit- 
tingly characterize  O.  biennis  and  0.  lamarckiana,  and  as  the  descriptions  were 


6        MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF    THE    OENOTHERAS. 

made  before  0.  grandi flora,  the  only  known  species  which  might  be  confused 
with  0.  lamarckiana,  was  discovered,  the  inference  may  be  drawn  by  exclu- 
sion that  nothing  but  O.  lamarckiana  was  referred  to,  and  this  inference  is 
strengthened  by  the  fact  that  0.  lamarckiana  was  recognized  and  cultivated 
in  the  Paris  gardens  a  few  years  later. 

Of  the  cultures  that  have  been  carried  out  from  material  obtained  from 
various  sources,  the  following  are  selected  as  of  unusual  interest : 

Two  sheets  of  dried  material  of  0.  lamarckiana,  collected  by  Mr.  E.  P.  Bick- 
nell  at  Nantucket  City,  were  received  in  September,  1904.  These  plants  had 
been  found  in  the  previous  month,  growing  near  a  cottage,  and  had  spread  to 
an  adjoining  waste  lot.  A  visit  was  made  to  the  place  in  August,  1905,  by 
the  senior  author,  but  no  trace  of  the  plant  could  be  seen,  although  it  was 
found  again  in  1906  by  Mr.  Bicknell.  This  journey  was  inspired  by  the  behav- 
ior of  the  seeds  taken  from  the  herbarium  specimen  and  sown  in  sterilized  soil 
in  November,  1904.  No  particular  attention  v/as  paid  to  the  lot,  but  on 
January  27,  1904,  24  plantlets  representing  the  widest  diversity  observable 
were  transplanted  to  small  pots  in  accordance  with  the  usual  practice.  Six  of 
these  corresponded  quite  exactly  to  the  mutant  0.  albida.  Four  of  these 
died  before  May,  1905,  at  which  time  the  remainder  were  transplanted  to  the 
experimental  grounds.  All  of  the  other  individuals  developed  in  accordance 
with  qualities  of  0.  lamarckiana,  with  a  maximum  amount  of  color  in  the  buds, 
and  also  a  maximum  number  of  basal  branches  of  some  length.  The  pale 
mutants,  however,  did  not  proceed  beyond  the  rosette  stage  in  1905. 

The  owner  of  the  grounds  in  which  the  original  plants  were  growing  in 
Nantucket  could  not  give  any  information  as  to  the  origin  of  the  culture, 
except  to  say  that  it  had  been  started  from  seeds  a  great  many  years  before. 

A  lot  of  seeds  taken  from  a  trade  packet  sold  by  Vilmorin  &  Company,  of 
Paris,  were  sown  in  sterilized  soil  on  Januar}^  3, 1905.  The  record  shows  that 
192  duplicates  of  the  parental  type  had  been  discarded  when  the  1 1  remaining 
ones  were  transplanted  to  the  experimental  garden  on  May  16.  Of  these  but 
3  were  mutants,  including  i  lata,  i  nanella,  and  i  albida.  The  mutants  were 
thus  seen  to  constitute  but  1.5  per  cent  of  the  total  culture. 

Far  the  most  interesting  material  and  records  were  obtained  from  England, 
and  it  was  found  that  0.  lamarckiana  has  been  growing  in  some  profusion 
in  certain  localities  in  that  country  for  several  years. 

In  1905  Mr.  H.  Stuart  Thompson  (Thompson,  1905)  called  attention  to 
Oenothera  biennis  L.  and  0.  odorata  Jacq.,  growing  on  sandhills  in  Lancashire 
and  Somersetshire,  and  with  the  view  of  comparing  the  native  North  American 
0.  biennis  with  the  plant  so  called  growing  in  England,  Mr.  C.  Theodore  Green 
was  applied  to  for  seeds  of  the  plant  growing  in  Liverpool  district.  To  him, 
therefore,  we  are  indebted  for  seeds,  a  photograph  of  the  Lancashire  0.  biennis, 
and  the  note  on  the  following  page  (Green,  1902). 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF    THE    OENOTHERAS.        7 

Oenothera  "biennis"  Linn^us. 

Form  as  found  on  both  sides  of  the  River  Mersey,  from  the  estuary  of  the  River  Dee  to 
Southport,  1 8  miles  north  of  Liverpool.  It  is  especially  luxuriant  about  Formby  Sandhills, 
9  miles  north  of  Liverpool,  and  about  Bidston  railway  junction,  3  miles  north  of  Birken- 
head. It  flourishes  in  sandy  waste  ground,  chiefly  among  the  hills  of  blown  sand  along  the 
coast,  and  extends  a  few  miles  inland  in  scattered  groups  of  plants.  It  also  is  found  fre- 
quently in  cultivated  ground  in  cottage  and  other  gardens  from  the  Dee  to  Southport.  I 
do  not  know  the  date  of  its  first  appearance  in  the  Liverpool  district,  but  it  was  mentioned 
in  Dickinson's  Flora  of  Liverpool  publication,  1851  and  1855. 

F.  W.  Webb  saw  it  near  Leasowe  (i860),  5  miles  southwest  of  Liverpool,  and  T.  Sansom 
at  New  Brighton  (Dick,  Flora). 

I  have  known  it  in  various  parts  of  this  peninsula  of  W'irral  (between  Mersey  and  Dee) 
since  1892. 

It  has  in  the  last  five  years  extended  greatly  about  Bidston  Junction  among  the  sand . 
There  is  much  more  of  it  north  of  Liverpool.  In  C.  C.  Babington's  Flora  it  is  given  as 
"American"  and  its  home  "Lancashire  Sandhills."  When  newly  opened  the  flowers  are 
very  fragrant,  but  only  last  twenty-four  hours. 

Seeds  inclosed,  more  of  which  I  can  send  if  desired;  came  from  Bidston  Junction.  This 
plant  is  identical  with  that  of  North  Liverpool. 

At  Bidston,  the  characteristic  sandhill  plants  among  which  it  flourishes  are  Erysimum 
chetraiithiodcs,  Brassica  vionensis,  Anthyllis  vulncraria,  Psamma  arcnaria,  Equisetum  max- 
imiim,  Lychnis  alba,  Senccio  jacobaea,  Salix  arenaria. 

It  extends  chiefly  along  the  railway  lines,  which  is  suggestive  as  to  its  origin. 

Inclosed  enlargement  from  one-fourth  plate  shows  the  plant  in  situ,  with  flowers  and 
fruit  stems  in  end,  September,  1905.  This  year  I  saw  the  same  type  of  this  plant  about  the 
railway  at  Llangollen  in  the  northwest. 

^Ir.  H.  Stuart  Thompson  (Thompson,  1906)  quotes  Watson's  New  Botan- 
ist's Guide  (1837)  as  the  first  record  of  0.  biennis  on  the  coast  of  Somerset  and 
says,  "indeed  it  is  quite  a  feature  in  the  landscape."  It  was  this  sentence 
that  aroused  interest  in  these  special  plants,  as  anyone  conversant  with  the 
weed-like  aspect  of  the  0.  biennis  of  this  region  would  scarcely  credit  them 
with  anything  like  a  spectacular  appearance. 

Hall  in  1845  (Hall,  1845,  p.  37)  speaks  of  the  plant  as  being  extensively 
naturalized  on  the  Liverpool  sandhills,  and  James  Edward  Smith  in  1806 
(Smith,  1806,  p.  1534)  figures  a  plant  that  was  gathered  on  the  extensive  and 
dreary  sandbanks  on  the  coast  a  few  miles  north  of  Liverpool,  where  millions 
of  the  same  species  have  been  observed  by  Dr.  Bostock  and  ]\Ir.  John  Shep- 
herd, perfectly  wild,  covering  a  large  tract  between  the  first  and  second  range 
of  sandhills.  Some  natural  cause  has  no  doubt  established  it  here,  though 
possibly  from  the  opposite  side  of  the  Atlantic.  The  Smith  plate  and  the 
plant  figured  by  Baxter  (Baxter,  1839,  p.  257)  certainly  do  not  represent  the 
common  North  American  0.  biennis,  and  the  closing  note  in  Baxter,  referring 
to  the  sudden  expansion  of  the  flowers,  applies  more  readily  to  such  large- 
flowered  species  as  0.  grandiflora,0.  lamarckiana,  and  O.argillicola  than  to  the 
smaller-flowered  ones,  like  0.  biennis,  0.  muricata,  and  0.  oakesiana. 


8        MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF   THE    OENOTHERAS. 

A  recent  writer  on  ths  flora  of  Lancashire  records  that  0.  biennis  has  been 
estabhshed  there  for  the  last  70  or  80  years  and  that  "whenever  the  land  is 
disturbed,  or  the  sand  removed  to  form  new  roads,  this  plant  is  one  of  the 
earliest  to  grow  upon  it,  and  although  its  conspicuous  flowers  make  it  an  easv 
prey  for  constant  plucking,  it  survives  these  depredations  and  continues  to 
spread  more  and  more." 

\\'ith  Mr.  Green's  photograph  and  these  references  in  mind,  it  was  with  no 
little  curiosity  that  the  cultures  from  his  seeds  were  watched.  The  seed- 
lings raised  from  them  earlv  in  1906  proved  to  be  indubitable  O.  lamarckiana. 
In  the  seed-pan  2  seedlings  of  0.  lata  and  4  seedlings  of  0.  rubn'ncrvis  were  also 
recognized  among  a  preponderating  number  of  the  O.  lamarckiana.  These 
plants  were  observed  unto  maturity.  0.  rubrinervis  displayed  all  the  char- 
acters ascribed  to  it  by  Professor  De  Vries,  with  the  exception  that  they  were 
not  quite  as  large  as  some  that  had  previously  been  studied  in  the  New  York 
Botanical  Garden.  The  two  plants  of  0.  lata,  on  the  contrary,  were  more 
robust  than  any  we  had  had  in  New  York  from  Professor  De  Vries's  seed,  and 
the  pollen  being  more  abundant  than  is  usual  in  the  species,  efforts  were  made 
to  self-pollinate  the  flowers  in  the  attempt  to  obtain  pure  seed.  In  due  time 
a  few  capsules  ripened,  containing  a  very  small  amount  of  seed,  from  which 
were  raised  in  December,  1906,  8  0.  lamarckiana,  10  0.  lata,  2  0.  oblonqa,  and 
1  0.  albida  seedlings. 

Before  these  facts  were  ascertained,  the  presumption  was  that  the  British 
O.  biennis  was  a  slightly  larger-flowered  evening-primrose,  and  in  our  cul- 
tures from  foreign  seed  we  usually  named  it  "European  biennis,"  of  which 
no  real  equivalent  appears  to  be  known  here.  Whether  our  native  small- 
flowered  species  is  also  known  in  Great  Britain  is  not  apparent  from  the  man- 
uals, as  the  species  there  recorded  indicates  a  more  ornamental  plant  than 
the  one  with  which  we  are  familiar  here,  and  ours  is  certainly  not  "adapted  to 
the  shrubbery." 

The  misunderstanding  in  regard  to  the  identity  and  provenience  of  O. 
biennis,  0.  lamarckiana,  and  0.  muricata  seems  to  be  vfidespread.  There  is 
now  no  doubt  as  to  the  identity  and  original  habitat  of  0.  grandiflora  Alton. 
The  time  and  manner  of  its  introduction  into  England  are  also  known.  It 
escaped  from  gardens  after  it  was  taken  into  England  in  1778,  and  is  now 
found  growing  wild  in  many  places,  according  to  unverified  reports. 

But  as  to  the  advent  of  lamarckiana  we  are  still  at  a  loss  for  positive  proof. 
The  plant  referred  to  by  Miller,  if  correctly  identified  with  this  form,  would 
place  it  in  Haarlem  in  1757,  21  years  before  the  discovery  of  grandiflora. 
The  next  we  hear  of  the  grandiflora  of  Lamarck  is  in  the  Paris  garden  in  1797, 
which  was  seen  by  Seringe  not  to  be  the  grandiflora  of  Alton  as  known  in 
England  and  was  renamed  after  its  illustrious  discoverer.  Next  we  have  the 
striking  form  which  is  described  above  as  appearing  on  the  coast  of  Somerset 


MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF    THE    OENOTHERAS.        9 

in  1837,  and  at  other  places  since  then,  forming  now  quite  a  feature  of  the 
Liverpool  sandhills,  which  has  been  shown  by  our  cultures  to  be  the  lamarck- 
iana  around  which  so  much  interest  clusters.  It  would  seem  difficult  in  the 
light  of  these  facts  to  find  any  basis  upon  which  the  conclusion  that  this  plant 
had  been  derived  from  0.  grandi flora  might  rest. 

The  plant  originally  introduced  into  Europe  under  the  name  of  0.  hicnnis 
seems  to  have  been  a  large-flowered  form  altogether  different  from  the  plant 
under  that  name  as  known  in  America.  Then  O.  lamarckiana  grown  from 
imported  seeds  or  their  progeny,  and  O.  qrandiflora  from  Alabama,  present 
a  very  diflferent  aspect  and  have  but  little  resemblance  to  the  0.  biennis  of 
the  waste  lands  of  eastern  America. 

THE  MUTABILITY  OF  OENOTHER.4  LAMARCKIANA. 

About  2500  plantlets  of  0.  lamarckiana  derived  from  various  sources  have 
been  brought  under  observation,  as  described  above.  Of  these  106  or  a  little 
over  4  per  cent  were  mutants.  One  culture  (C.1.2)  appeared  to  give  a  much 
higher  proportion  of  mutants,  but  a  failure  to  make  a  record  of  one  lot  of 
discarded  plants  makes  it  impossible  to  confirm  this.  The  behavior  of  this 
lot  of  plants  and  also  the  confusion  of  fasciated  specimens  of  0.  lamarckiana, 
in  the  rosette  stage,  with  mutants,  led  the  senior  author  to  assert  that  the  pro- 
portion of  mutant  derivatives  in  pedigreed  cultures  of  O.  lamarckiana  had 
been  increased  from  5  to  6  per  cent  of  the  progeny  (MacDougal,  1905). 
In  no  culture  of  later  date  has  it  been  possible  to  exceed  the  maximum  propor- 
tion of  5  per  cent,  and  the  greater  majority  fall  much  below  this  figure.  While 
the  coefficient  of  mutability  has  not  been  modified  yet,  one  or  two  forms  have 
appeared  which  were  not  recognizable  in  a  rosette  stage  by  Professor  De 
Vries,  and  probably  constitute  an  extension  of  the  range  of  mutability.  This, 
however,  may  not  be  interpreted  to  mean  an  effect  of  local  conditions.  All 
of  the  mutants  now  known  did  not  occur  in  the  cultures  in  Amsterdam 
during  the  first  few  years  of  the  cultures.  Many,  in  fact,  occur  so  rarely 
that  by  the  law  of  probability  it  is  necessary  to  grow  many  thousands  of  speci- 
mens in  order  to  secure  one  individual  of  the  rare  type.  In  accordance  wnth 
this  same  principle  it  is  to  be  seen  that  the  spreading  of  the  cultures  of  O. 
lamarckiana  may  be  expected  to  bring  to  light  other  rarer  derivatives.  When 
such  forms  are  found  it  is  not  to  be  interpreted  as  a  result  of  local  conditions 
of  the  culture  in  which  they  appear,  but  simply  in  accordance  wnth  the  coeffi- 
cient of  mutability  of  the  parental  type  with,  respect  to  the  form  in  question. 

The  uniformly  low  frequency  of  mutants  in  material  procured  from  seeds- 
men is  a  matter  not  altogether  understood.  Among  the  probabilities  to  be 
taken  into  account  is  that  the  seeds  may  have  lain  in  storage  for  more  than 
one  season  before  being  planted,  in  consequence  of  which  the  unequal  hardi- 
ness of  the  mutants  and  the  parent  might  operate  to  decrease  the  apparent 


lO     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 

frequency  of  the  derivatives.  No  proof  has  been  produced  to  show  that  the 
frequency  of  mutations  might  be  increased  beyond  the  Hmit  of  5  per  cent 
found  by  De  Vries,  but  it  is  a  well-known  fact  that  mutations  of  all  kinds  may 
be  decreased  by  inadequate  nutrition,  and  it  is  quite  possible  that  the  parental 
plants  grown  to  furnish  seeds  did  not  have  the  care  and  nourishment  of  pedi- 
greed individuals  used  in  the  cultures.  The  appearance  of  the  rare  ruhrinervis 
in  the  culture  might  be  due  to  a  hybridization  with  that  form,  since  no  evi- 
dence has  been  offered  to  show  that  the  fertilizations  were  guarded. 

The  five  cultures  described  together  furnished  albida,  nanella,  oblonga,  lata, 
gigas,  scintillans,  and  nine  other  forms  not  positively  identifiable  at  this  time. 
Seeds  have  been  preserved,  however,  and  it  is  hoped  we  may  be  able  to 
present  descriptions  of  at  least  some  of  the  forms  at  the  close  of  another  year. 

Since  these  cultures  were  carried  through,  De  Vries  has  published  the 
results  of  his  cultures  from  seeds  obtained  from  seed  merchants.  2000  plantlets 
from  seeds  obtained  from  Haage  &  Schmidt  of  Erfurt  contained  i  specimen 
of  ruhrinervis,  which  arises  but  rarely  in  any  culture,  i  of  oblonga,  and  3  of 
nanella. 

Two  cultures  were  made  from  seeds  furnished  by  Vilmorin  and  grown  in 
1 898- 1 899.  The  first  gave  14  nanella,  3  lata,  3  scintillans,  1  albida,  i  oblonga, 
and  a  few  other  divergent  forms  in  3500  seedlings.  A  second  test  yielded 
3  lata,  I  nanella,  and  i  ruhrinervis  (?)  in  600  seedlings.  In  both  cases  the  pro- 
portion of  the  mutants  was  not  above  5  per  cent,  and  was  even  less  than  that 
described  above  (De  Vries,  1905). 

In  view  of  the  numerous  observations  described  above  upon  material  from 
the  most  widely  separated  sources,  which  show  mutability,  suggestions  that 
the  mutability  of  Lamarck's  evening-primrose  has  been  consequent  upon 
its  segregation  in  the  Amsterdam  Botanical  Garden  may  no  longer  be  taken 
seriously,  although  no  doubt  the  well-worn  phrase  will  be  duly  rehearsed 
from  time  to  time  by  careless  critics.  0.  lamarckiana  may  be  of  hybrid  origin, 
and  what  species  may  not  be;  but  if  so  this  origin  is  not  recent,  nor  can 
sufiicient  time  be  found  to  hunt  down  all  of  the  unprofitable  alternatives  that 
are  offered  for  consideration. 

Exact  records  have  already  been  made  of  lamarckiana,  ruhrinervis,  nanella, 
and  gigas  in  previous  papers  (MacDougal,  Vail,  Shull  &  Small,  1903  and 
1904),  and  below  are  the  principal  taxonomic  characters  of  albida,  oblonga, 
scintillans,  hrevistylis,  and  lata,  by  which  systematic  descriptions  of  9  of  the 
15  mutant  derivatives  iteratively  yielded  by  lamarckiana  are  afforded  from 
material  grown  in  New  York  (see  plate  i). 

Note. — The  illustrations  of  0.  lamarckiana  and  O.  ruhrinervis  prcvionsly  published  by 
MacDougal  (1903)  were  reduced  without  proper  indication  of  the  scale. 

The  capsule  of  0.  gigas  is  incorrectly  given  in  another  paper  as  2  nun.  in  length  when  it 
should  be  2  cm.     (MacDougal,  Vail,  Shull  &  Small,  1905.) 


PLATE  2. 


^=^^ 


Oenothera  scinlillans.      1,  leaf  from  rosette;  2,  leaf  from  stem;  3,  bract;  4,  bud  with 
bract;  5,  flower  with  petals  removed;  6,  petal;   7,  capsule. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF    THE)    OENOTHERAS.      II 

OENOTHERA  SCINTILLANS. 

O.  scintiUans  was  first  seen  to  originate  as  a  mutant  in  1888  and  has  appeared 
in  this  manner  in  14  individuals  at  different  times  in  De  Vries's  cultures.  As 
previously  described  in  this  paper,  it  has  appeared  in  4  specimens  in  the  New- 
York  Botanical  Garden  during  the  present  year. 

O.  scintiUans  is  to  be  noted  as  an  ever-sporting  form  which  in  some  strains 
gives  a  progeny  60 or  70  per  cent  of  which  is  composed  of  lata,  oblonga,  lamarck- 
iana,  and  nanella,  while  in  other  strains  but  30  per  cent  will  be  included  in 
these  forms.  The  forms  thus  derived  from  scintiUans  are  in  no  wise  different 
in  their  hereditary  qualities  from  the  same  types  derived  from  other  and  pure 
lineages.  Cultures  were  made  from  seeds  furnished  by  Professor  De  Vries 
from  a  strain  which  was  supposed  to  yield  60  to  70  per  cent  of  the  other  types 
named. 

Seventy-eight  individuals  were  obtained  from  the  culture,  15  of  which  resem- 
bled scintiUans,  ^6  lamarckiana,  16  oblonga,  and  i  was  present  which  did  not 
reach  maturity,  but  suggested  some  of  the  incomplete  forms  mentioned  in  ' '  Die 
Mutationstheorie,"  ^rohahXy  sublinearis.  In  the  seedlings  3  months  old,  the 
rosettes  were  14  to  17  cm.  in  diameter,  dense,  and  spreading  flat  on  the  ground ; 
outer  leaves,  6  to  7  cm.  long,  blades  about  5  cm.  long,  2.5  to"  3  cm.  wide, 
oblong-obovate,  widest  about  the  middle,  obtuse  or  acutish  at  the  apex, 
tapering  gradually  to  the  white-margined  petiole,  blue-green,  shining,  puber- 
ulent,  with  occasional  small  red  spots  above,  glabrous  or  nearly  so,  and  lighter 
green  beneath,  somewhat  brittle.  At  6  months  of  age  the  rosettes  were 
dense,  the  leaves  lanceolate,  obscurely  and  minutely  denticulate,  green  and 
shining,  puberulent  on  margin  and  sparingly  on  upper  surface,  midvein  broad, 
petioles  variously  winged. 

The  adult  plant  was  short,  with  few  branches  which  were  fairly  rigid  and 
ascending.  The  lower  part  of  the  stem  appeared  to  be  terete  or  nearly  so, 
while  the  upper  portion  was  angled  and  channeled.  The  stem  and  branches 
were  hirsute  with  spreading  hairs. 

The  stem-leaves  were  finely  pubescent  on  both  surfaces  and  m_uch  crinkled 
on  the  margins  and  midribs;  irregularly  toothed,  oblong,  7 to  9  cm.  long,  30  to 
35  mm.  wide,  abruptly  tapering  at  base,  acutish  at  the  apex.  The  petioles 
were  short  and  the  entire  organ  was  a  deep  shining  green. 

Conic  portion  of  the  bud  15  to  20  mm.  long,  finely  pubescent,  with  short, 
spreading  hairs.  Free  tips  5  mm.  long,  erect.  Hypanthium  30  mm.  long,  and 
reflexed  calyx-lobes  25  cm.  long.     Ovary  6  to  7  mm.  long. 

The  petals  are  thin,  25  mm.  long  and  30  mm.  in  width,  barely  emarginate. 
Filaments  15  to  18  mm.  long,  very  slender  and  strongly  upcurved.  Anthers 
7  to  8  mm.  long.  Pistils  slightly  longer  than  the  stamens ;  lobes  of  the  stigma 
very  slender,  4  to  5  mm.  long  (plate  2). 


12      MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF   THE    OENOTHERAS. 

OENOTHERA  BREVISTYLIS. 

A  culture  was  made  from  seed  harvested  by  De  Vries  in  Amsterdam  in  1 899 
and  said  by  him  to  contain  about  25  per  cent  hrevistylis,  the  seed  being  the 
product  of  the  first  generation  of  a  cross  between  0.  lamarckiana  and  0. 
bvevistylis.  Twenty-one  plantlets  appeared  and  were  inspected,  and  some 
seemed  to  form  rosettes  with  much  more  rounded  and  abruptly  pointed  apices 
than  0.  lamarckiana.  This  was  the  only  distinction  that  could  be  made 
between  the  parental  type  and  the  derivative,  during  what  would  naturally 
be  the  first  year  of  development.  In  addition  one  example  of  0.  nanella  as  a 
mutant  from  the  parental  type  was  included.  With  the  beginning  of  the 
formation  of  lengthened  internodcs  in  the  stem  but  little  difference  between 
the  plants  was  discernible  until  flower-buds  appeared.  At  that  time  the 
leaves  on  the  terminal  portions  of  the  branches  and  the  bracts  seemed  rela- 
tively much  broader,  and  the  flower-buds  might  be  distinguished  at  some 
distance  by  the  fact  that  they  were  more  cylindrical  than  the  parental  type, 
and  were  abruptly  short-pointed.  An  examination  of  the  structure  of  the 
flower  revealed  the  fact  that  the  style  is  much  shorter  than  in  0.  lamarckiana 
and  that  the  stigma  generally  appears  in  the  throat  of  the  flower  and  some- 
times is  not  to  be  seen  without  tearing  the  hypanthium  apart. 

It  has  been  noted  by  De  Vries  that  the  length  of  the  style  varies  widely 
(as  much  as  i  cm.),  and  in  this  is  offered  an  example  confirmatory  of  results 
communicated  by  the  authors  in  a  recent  paper  (MacDougal,  Vail,  Shull  & 
Small,  1905),  in  which  the  mutant  characters  were  found  to  offer  a  wider 
amplitude  of  variability  than  the  correspondent  characters  of  the  parental 
type.  In  addition  to  the  above  general  characters,  it  is  to  be  noted  that,  as  a 
consequence  of  the  extreme  shortness  of  the  style,  pollination  fails  in  many 
cases  and  comparatively  few  capsules  are  matured. 

The  evidence  offered  by  this  plant  is  of  the  greatest  interest  in  connection 
with  questions  concerning  the  survival  of  mutants  in  the  habitat  of  the  paren- 
tal form,  especially  in  view  of  the  intemperate  indulgence  which  many  authors 
show  in  theoretical  discussions  upon  questions  of  this  character. 

0.  hrevistylis  was  discovered  August  20,  18S6,  by  De  Vries,  as  represented 
by  2  individuals.  Despite  the  greatest  care  and  the  most  rigid  inspection  it 
has  never  been  observed  to  arise  in  any  of  the  cultures  in  Amsterdam  or  New 
York,  and  the  supposition  is  certainly  allowable  that  it  is  no  longer  included 
among  the  nmtants  given  off  by  the  parental  type.  The  characteristic  qual- 
ities of  O.  hrevistylis  are  recessive  when  hybridized  with  0.  lamarckiana,  and 
consequently  the  first  generation  resembles  O.  lamarckiana  in  outward  form 
but  carries  0.  hrevistylis,  which  reappears  in  the  next  or  the  second  generation 
of  the  hybrid,  forming  on  an  average  one-fourth  of  the  progeny,  according  to 
the  simple  Mendelian  formula.     0.  hrevistylis  is  a  retrogressive  departure  from 


PLATE  3. 


Oenothera  breris/ylis.  1,  leaf  from  mature  rosette;  2,  bract  and  capsule;  3,  stem-leaf; 
4,  flower  with  petals  removed;  5,  bud;  6,  hypanthium  split  open,  showing  the  short, 
imperfect  style;  7,  petal;  8.    bract;  9,  capsule. 


PLATE   4. 


2  4 

Oenothera  lata.      1,  leaf  from   mature    rosette;    2,   bract   and  capsule;   3,   bract;  4,   bud  and   bract; 

5,    flower  with  petals  taken  off;  6,  petal. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS.      1 3 

the  parental  type,  matures  comparatively  few  seeds,  and  is  dominated  by  the 
parental  characteristics  in  crossing,  and  it  has  survived  in  its  original  habitat 
and  blooming  specimens  have  been  observed  there  for  twenty  years  in  com- 
petition with  the  parental  form. 

The  more  important  anatomical  features  of  O.  brevistylis  were  described 
by  Pohl  (1895),  but  it  seems  desirable  that  its  general  taxonomic  aspect,  as 
presented  in  tha  cultures  in  New  York,  should  be  put  on  record. 

The  rosettes  are  open,  leaves  ovate-spatulate,  crinkled,  the  upper  ones 
approximitely  denticulate,  bright,  dark  green,  puberulent  beneath,  less  so 
above,  conspicuous  broad  veins,  petiole  winged  to  near  the  base,  much  more 
obtuse  and  rounded  at  apex  than  0.  lamarckiana. 

The  adult  shoot  resembles  0.  lamarckiana  very  closely,  even  in  the  form 
of  the  stem-leaves.  The  bracts,  however,  seem  slightly  thinner  and  appear 
somewhat  broader  than  those  of  the  parental  type. 

The  conic  parts  of  the  buds  are  2  to  5  cm.  long,  appearing  more  cylindrical 
than  the  parental  form,  and  are  covered  with  very  short,  spreading  hairs. 
The  buds  are  distinctly  tinged  w4th  red,  and  the  free  erect  tips  are  unequal 
and  are  not  spreading. 

The  petals  are  45  to  50  mm.  long  and  40  mm.  wide,  being  firm  and  more  or 
less  deeply  emarginate.  The  filaments  are  2  cm.  long,  the  anthers  i  cm.  long 
and  slender.  The  pistil  does  not  usually  emerge  from  the  tube  of  the  hypan- 
thium,  the  stigma  being  seen  in  the  very  throat  of  the  flower.  The  stigmatic 
lobes  are  very  irregularly  developed. 

But  few  capsules  are  matured.  The  capsules  are  15  to  18  mm.  long  and 
6  to  7  mm.  in  diameter  at  the  base,  being  finely  pubescent  with  a  few  spreading 
hairs,  bright  green  and  slender,  tapering  slightly  toward  the  apex. 

The  hypanthium  in  this  species  bears  about  the  same  relation  to  the  reflexed 
sepals  as  in  the  parental  form  (plate  3). 

OENOTHERA  LATA. 

0.  lata  was  first  seen  by  De  Vries  in  the  plantlets  produced  by  seeds  har- 
vested at  Hilversum,  Holland,  in  1886,  and  brought  into  bloom  in  1887,  and 
he  has  observed  the  origin  of  a  total  number  of  493  as  mutants  in  a  progeny  of 
the  parental  type  embracing  130,000  plants. 

It  has  been  observed  to  arise  in  New  York  from  the  third  generation  of 
a  strain  grown  from  seeds  furnished  by  Professor  De  Vries,  and  also  in  seeds 
obtained  from  Vilmorin  in  Paris. 

A  sowing  was  made  of  seeds  of  0.  lata  X  0.  lamarckiana,  furnished  by  Pro- 
fessor De  Vries.  This  form  is  characterized  by  atrophied  stamens  which 
produce  only  a  few  pollen  grains,  of  fairly  normal  structure,  which  seem  to 
be  incapable  of  producing  fertilization;  but  when  the  plant  is  pollinated  by 
O.  lamarckiana,  lata  characters  appear  in  the  first  generation  as  forming  from 


14     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 

4  to  45  per  cent  of  the  progeny,  or  an  average  of  22  per  cent.  In  the  cultures 
made  in  New  York,  lata,  if  present,  was  mistaken  for  lamarckiana  in  the  seed- 
hng  stage  and  was  discarded.  Among  the  plants  of  this  progeny  which  were 
finally  brought  to  bloom  were  albida,  oblonga,  nanella,  and  another  type  not 
recognizable  by  the  authors. 

Laia  is  to  be  regarded  as  a  retrogressive  departure  from  the  parental  type, 
and  is  most  readily  recognized  by  the  rounded  leaves  of  the  rosettes,  and  these 
organs  become  very  much  crinkled  in  later  stages  of  development.  The 
leaves  are  of  a  deep  green,  and  the  flower-buds  are  much  thicker  than  any 
other  evening- primrose  known,  except  gigas,  perhaps. 

The  principal  taxonomic  characters  are  as  follows:  The  general  habit  of 
this  species  resembles  that  of  0.  gigas  more  nearly  than  any  other  of  the 
mutants,  being  sparingly  branched,  and  having  short,  stoutish  branches  on 
the  upper  portion  of  the  stem.  All  of  the  branches  are  ascending,  and  in 
many  of  the  plants  the  terminal  portion  of  the  stem  is  bent  over. 

The  stem  is  channeled  and  angled,  brittle,  and  both  stems  and  branches 
are  hirsute.  The  stem-leaves  are  finely  pubescent,  10  to  15  cm.  long,  4  to  5 
cm.  wide,  numerous,  and  with  the  laminae  remotely  and  shallowlv  toothed. 
The  leaves  of  the  rosettes,  even  in  the  very  early  stages,  are  noticeablv  rounded 
and  obtuse  at  the  apex,  and  those  of  the  lower  part  of  the  stem  are  spatulate- 
oblong  and  vary  to  ovate-oblong  on  the  upper  portions  of  the  stem.  Those 
on  the  upper  part  of  the  stem  are  either  obtuse  or  acutish  at  the  apex  and 
taper  to  a  margined  petiole  at  the  base.  The  leaves,  even  in  voung  rosettes, 
are  closely  bunched  into  a  head  and  are  so  heavily  crinkled  and  thrown  into 
convexities  and  concavities  between  the  veins  as  to  be  easily  distinguishable 
from  all  other  forms  in  all  stages  of  development. 

The  bracts  are  large,  ovate  to  ovate-oblong,  acute  or  obtuse  at  the  apex, 
cordate  or  subcordate,  and  clasping  at  the  base. 

The  conic  portions  of  the  heavy  buds  are  2  cm.  long  and  i  cm.  in  diameter  in 
the  basal  portion,  tapsring  so  slightly  as  to  appear  nearly  cylindrical,  and  are 
finely  pubescent  with  spreading  hairs.  The  free  tips  of  the  buds  are  spreading, 
stout,  4  to  5  mm.  long.  The  hypanthium  is  about  35  mm.  long,  stout,  finely 
pubescent  and  much  longer  than  the  reflexed  sepals  (plate  4  and  plate  5,  a). 

The  petals  are  thick,  crinkled,  and  do  not  expand  fully,  being  about  35  to  40 
mm.  long,  and  45  to  50  mm.  wide.  The  filaments  are  18  to  20  mm.  long, 
while  the  anthers  are  imperfect,  being  very  slender  and  about  8  mm.  in  length. 
The  microscopical  examination  of  the  anthers  showed  a  few  pollen  grains 
apparently  perfect,  yet  no  fertilization  has  ever  been  accomplished  with  them 
in  Amsterdam. 

In  a  recent  paper  R.  R.  Gates  (1907)  describes  the  results  of  an  investigation 
of  the  pollen  development  of  0.  lata  in  which  he  ascribes  the  failure  of  the 


PLATE  5. 


#       # 


t^ 


B 

A.     KosL-ttu  otOciiotlKia  l:il;i,  fr)iir  months  old,  anil  separate  leaves  of  the  same  age. 
H.      Rosette  o(  Oenothera  alhida.  tour  months  old,  and  sejiarate  leaves  of  the  same  age. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS.      1 5 

pollen  to  other  causes  than  the  ingrowth  of  the  tapetum  to  fill  the  loculus. 
He  found  that  development  may  proceed  to  the  formation  of  tetrads,  but 
frequently  degeneration  begins  in  the  resting  stage  of  the  first  mitosis.  The 
successful  pollinations  recorded  below,  however,  demonstrate  the  functional 
maturitv  of  the  pollen  in  many  cases.  Heterochromosomes  were  found  to 
arise  in  the  prophase  after  synapsis,  both  in  lata  and  in  the  lamarckiana  hybrid 
with  lata.  Inferential  support  was  secured  for  the  supposition  that  the 
changes  constituting  mutation  occur  during  the  reduction  divisions. 

The  pistil  is  barely  longer  than  the  stamens,  with  the  stigmatic  lobes,  heavy 
and  club-shaped,  6  to  7  mm.  long.  The  capsules  are  13  mm.  long,  about  6  to  7 
mm.  in  diameter  in  the  thickest  portion,  finely  pubescent,  many  angled,  and 
tapering  slightly  toward  the  apex.  The  terminal  rosettes  on  the  stem  and 
branches  were  small  and  regular. 

While  this  form  has  hitherto  been  found  incapable  of  producing  mature 
pollen  capable  of  effecting  fertilization  in  repeated  trials  made  in  Amsterdam 
and  New  York,  a  notable  exception  is  to  be  recorded.  A  package  of  seeds  of 
O.  lamarckiana,  which  is  growing  wild  near  Birkenhead,  England,  was  received 
from  Mr.  C.  Theodore  Green  in  1905.  A  sowing  made  from  this  lot  yielded  the 
parental  form,  rubrinervis,  and  a  few  individuals  of  lata,  which  agreed  with 
this  form  as  derived  directly  from  pure  cultures  so  far  as  all  external  charac- 
ters were  concerned.  Many  individuals  previously  examined  in  various 
laboratories  showed  pollen  which  appeared  normal  under  the  microscope,  yet 
no  fertilizations  could  be  secured.  In  the  present  instance,  however,  pollen 
appeared  so  plentiful  that  another  attempt  was  made,  the  treated  pistils 
being  carefully  guarded  from  pollination  from  the  parental  or  other  forms. 
A  few  pods  were  matured  and  some  seeds  were  secured  as  a  result.  These 
were  sown  as  soon  as  ripe,  in  September,  1906.  The  progeny  showed  10  lata, 
80  lamarckiana,  i  albida,  and  3  ohlonga.  No  material  difference  between  this 
and  progenies  resulting  from  pollination  with  lamarckiana  was  thus  shown. 
In  the  latter  case  the  proportion  of  lata  is  often  as  low  as  4  per  cent  of  the 
entire  prog^nv  and  has  not  been  recorded  to  have  gone  beyond  45  per  cent. 

OENOTHERA  OBLONGA. 

O.  ohlonga  was  first  observed  by  De  Vries  at  Amsterdam  in  1895,  although 
it  doubtless  appeared  in  his  cultures  previous  to  that  time,  as  he  suggests.  In 
all  it  has  arrived  in  700  mutants  and  in  the  pedigree-cultures  described  in  this 
paper  in  35  individuals,  only  a  part  of  which  came  into  bloom  in  August,  1905, 
agreeing  with  the  observations  in  Amsterdam.  Oblonga  is  one  of  the  most 
easily  recognizable  of  the  derivatives  of  0.  lamarckiana,  although  the  distin- 
guishing characters  do  not  readily  lend  themselves  to  taxonomic  description. 

The  rosettes  are  not  very  dense ;  the  leaves  in  the  young  rosettes  are  nar- 
rowly ovate-lanceolate,  rather  thick  and  fleshy,  with  broad  midveins  which 


1 6     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

are  sometimes  distinctly  reddish.  The  laminae  are  shining  green  above,  more 
or  less  puberulent  on  both  surfaces,  and  obscurely  denticulate. 

The  adult  plant  is  less  than  i  meter  in  height  and  is  very  sparingly  branched 
below,  with  a  few  very  short  branches  above.  The  basal  branches  are  ascend- 
ing and  do  not  reach  above  half  the  height  of  the  stem.  The  stems  are 
strongly  channeled  and  angled. 

The  stem-leaves  are  crowded,  hang  down,  and  are  pubescent  on  both  sur- 
faces, 7  to  lo  cm,  long,  oblong-elliptical,  remotely  and  shallowly  toothed, 
acutish  or  obtuse  at  the  apex,  and  irregularly  narrowed  into  a  short  margined 
petiole.  The  laminae  are  strongly  crinkled,  dark  green,  and  become  tinged 
with  crimson  with  age. 

The  bracts  are  oblong  or  ovate-oblong,  acute  at  the  apex,  cordate  or  sub- 
cordate  at  base.  The  buds  are  reddish,  the  conic  portion  being  about  2  cm. 
long  and  finely  pubescent  with  short  spreading  hairs  and  a  few  longer  ones. 
The  free  erect  tips  are  about  3  to  5  mm.  in  length.  The  petals  are  thin, 
generally  crinkled,  somewhat  emarginate  and  crenate,  and  are  3  cm.  long  and 
4  wide.  In  the  unfolding  of  the  flower  the  petals  open  only  so  far  as  to  form 
a  cup-shaped  corolla.  The  hypanthium  is  about  3  cm.  long,  slightly  longer 
than  the  calyx-lobes.  The  slender  ovary  is  about  8  mm.  long.  The  filaments 
are  15  mm.  long  and  the  anthers  8  mm.  The  pistils  are  slightly  longer  than 
the  stamens,  the  stigmatic  lobes  being  about  5  mm.  in  length. 

The  capsules  are  22  to  26  mm.  long  and  6  to  7  mm.  in  diameter  at  the  thick- 
est part,  finely  pubescent,  slightly  angled  and  shining  green,  tapering  slightly 
to  the  apex  (plate  6). 

OENOTHERA  ALBIDA. 

O.  albida  was  first  recognized  by  De  Vries  as  a  mutant  in  1888  and  was 
brought  into  bloom  in  1896.  It  is  distinguishable  even  in  the  earlier  stages 
by  the  paler  color  of  the  leaves,  which  in  the  rosettes  have  upturned  margins 
and  are  variously  twisted.  Seventeen  examples  appeared  in  the  cultures 
previously  described  during  1905,  of  which  6  were  mutants  from  0 .  lamarckiana 
as  found  introduced  on  Nantucket.  The  following  taxonomic  characters 
were  observed : 

Seedling  about  3  months  old:  Rosettes  15  to  17  cm.  in  diameter,  thin, 
somewhat  raised  above  the  slender  rootstock;  leaves  7  to  10  cm.  long,  2.5  to 
3  cm.  wide,  erect-spreading,  the  apex  recurved  and  nearly  touching  the 
ground;  blades  5  to  7cm.  long, oblong,  broadest  at  about  the  middle,  acutish 
or  mostly  obtuse  at  the  variously  rounded  apex,  gradually  tapering  into  the 
almost  translucent,  margined  petiole,  pale  light  green,  very  thick  and  brittle, 
margins  undulate-denticulate,  some  of  the  3^ounger  leaves  strongly  wrinkled 
on  the  margins.  About  2  months  later  the  rosettes  were  irregular,  leaves 
twisted  and  various-erected,  convexed  upwardly,  pale  yellowish-green,  approx- 


PLATE  6. 


Oenothera  oblcnga.      1,    leaves   from   mature    rosette;    7.  stem-leaf;    3,  bracts;    4,    petal;    5,    capsule; 

6,  bud;  7,  flower  with  petals  removed. 


PLATE  7 


Oenothera  albida.      1,  leaf  of  mature  rosette;  2,  stem-leaf;  3,   bract;  4,   bud;  5,   flower  with 

petals  removed;  6,  petal;  7,  capsule. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      1 7 

imately  denticulate,  oblong,  soft-hairy  on  both  surfaces,  petioles  winged  half- 
way to  base. 

The  adult  plant  was  seen  to  be  irregularly  branched,  the  branches  often 
exceeding  the  main  stem  in  length,  which  never  reached  a  height  of  a  meter. 
Stem  zigzag,  rather  stout,  channeled  and  angled,  branching  mostly  at  base,  both 
branches  and  stems  very  brittle.  Branches  and  stems  hirsute,  with  spreading 
hairs  and  closely  pubescent.     Internodes  short,  with  the  leaves  crowded. 

Stem-leaves  oblong  to  oblong-lanceolate,  more  or  less  regularly  denticulate, 
mostly  obtuss  or  acutish.  nearly  sessile,  tapering  at  base,  finely  pubescent 
on  both  surfaces,  very  brittle  and  somewhat  crinkled. 

The  bracts  are  oblong,  acutish,  or  obtuse,  slightly  clasping,  subcordate, 
erected  when  young,  reflexed  when  mature,  margins  upturned. 

The  buds  are  distinctly  reddish,  3  cm.  long,  finely  pubescent,  with  spreading 
hairs  and  the  free  tips  erect  and  6  mm.  long. 

Hypanthium  4  cm.  long,  slightly  longer  than  the  sepals.  Petals  thin, 
35  to  40  mm.  long,  and  40  to  50  mm.  wide,  reflexed  from  the  middle,  deeply 
emarginate  at  the  apex.  Filaments  2  to  3  cm.  long,  anthers  12  mm.  long. 
Pistils  longer  than  the  stamens,  with  the  stigmatic  lobes  5  mm.  in  length. 

Capsules  35  mm.  long  and  5  to  6  mm.  in  diameter  at  thickest  portion,  finely 
pubescent  and  reddish,  tapering  slightly  to  apex.  It  is  noticeable  that  com- 
paratively few  capsules  are  matured  by  this  species.  The  terminal  rosettes 
are  symmetrical  and  the  buds  erect  and  prominent  (plate  5,  b,  and  plate  7). 

HYBRIDIZATION  OF  A  MUTANT  AND  ITS  PARENTAL  TYPE. 

In  order  to  test  the  results  which  have  been  presented  by  Professor  De  Vries 
with  respect  to  the  behavior  of  mutants  in  hybridization,  a  number  of  flowers 
of  0.  lamarckiana  were  castrated  early  in  order  to  avoid  entirely  any  proba- 
bility of  self-pollination,  and  these  flowers  were  pollinated  from  unopened 
buds  of  0.  ruhrinervis.  The  resulting  capsules  were  harvested  early  in  Sep- 
tember, 1904,  and  sown  in  sterilized  soil  in  the  greenhouses  in  November,  1904. 
Early  in  January  the  seedlings  were  large  enough  to  make  the  separate  types 
discernible,  and  the  task  of  discarding  the  duplicates  was  begun.  The  entire 
progeny  was  found  to  include  90  of  lamarckiana  (and  one  of  its  mutants,  of 
which  I  was  present)  and  221  rubrinervis.  The  ruhrinervis,  which  is  to  be 
regarded  as  a  progressive  mutant  of  latnarckiana,  has  been  seen  to  surpass  it  in 
vegetative  vigor  and  rapidity  of  growth,  besides  producing  seeds  quite  as 
plentifully,  and  is  sometimes  dominant  when  crossed  with  the  parent.  Thus 
De  Vries  found  that  when  the  above  cross  was  made  he  obtained  19,  24,  68, 
and  74  per  cent  of  rubrinervis  in  four  different  lots,  giving  an  average  of 
totals  of  46  per  cent  rubrinervis  produced  in  such  crosses.  The  culture 
described  above  yielded  71  per  cent  rubrinervis. 


THE   FL^UCTUATIONS   OF  OENOTHERA   LAMARCKIANA   AND 

ITS  MUTANTS. 

By  George  Harrison  Shull. 

The  results  of  statistical  studies  made  in  1904  on  Oenothera  lamarckiana  Ser. 
and  two  of  its  mutants,  0.  nanella  and  O.  rubrinervis,  were  so  suggestive  that 
it  was  thought  desirable  to  make  a  further  attempt  to  trace  the  relations  of 
the  mutants  to  their  parental  form,  as  indicated  by  the  variations  in  some 
of  their  differentiating  characters.  In  order  to  reduce  the  probable  error,  it 
was  necassary  to  secure  data  from  a  larger  number  of  individuals  than  the 
cultures  at  the  New  York  Botanical  Garden  could  supply.  Consequently  cul- 
tures of  several  of  the  Oenotheras  were  begun  at  the  vStation  for  Experimental 
Evolution. 

The  cultures  which  form  the  basis  of  this  comparative  study  were  the 
following: 

(a)  About  90  spscimens  of  0.  lamarckiana  were  grown  from  seed  supplied 
under  the  designation  D.1.8  by  the  senior  author,  being  seed  from  plants 
guarded  and  purely  self-fertilized  for  three  generations  in  the  New  York 
Botanical  Garden,  and  thirteen  generations  preceding  in  the  Botanical  Gar- 
den at  Amsterdam.  In  this,  as  in  all  the  following  cultures,  no  selection  of 
specimens  was  made,  the  desired  number  of  plants  having  been  taken  consecu- 
tively, beginning  at  one  side  of  the  seed-pan ;  and  in  all  cases  the  arrangement 
in  the  garden  was  the  same  as  that  adopted  at  the  New  York  Botanical  Garden, 
i.  e.,  the  plants  were  placed  at  a  uniform  distance  of  i  meter  from  each  other. 
One  of  these  90  individuals  was  an  O.  lata  and  was  omitted  from  the  statistical 
studies  on  this  plot.     The  remaining  89  specimens  were  typical  0.  lamarckiana. 

(b)  Ten  rosettes  of  Oenothera  lamarckiana  were  received  by  mail  from  Prof. 
De  Vries  April  7,  1905,  having  been  collected  by  him  in  the  open  field  near 
Hilversum,  Holland,  where  over  twenty  years  ago  he  secured  the  original 
stock  for  his  pedigree-cultures;  9  of  these  came  to  bloom  and  were  uni- 
form in  appearance,  agreeing  well  with  the  other  cultures  of  O.  lamarckiana 
in  all  characters  but  those  whose  differences  could  be  appropriately  attributed 
to  the  fact  that  these  plants  had  grown  as  biennials,  while  the  others  had  been 
forced  to  annual  bloom  by  giving  an  early  start  in  the  propagating  house. 
The  chief  differences  between  these  plants  and  those  of  the  culture  described 
above  were  seen  in  the  greater  height  and  more  numerous  and  longer  branches 
of  the  plants  from  Hilversum.  The  rosette  leaves  were  more  numerous  and 
larger  in  both  dimensions,  but  had  the  same  form  and  the  same  degree  of 
crinkling. 

18 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      1 9 

(c)  Eighty-eight  specimens  were  grown  from  seed  of  an  O.  rubrinervis 
which  had  appeared  as  a  newly  arisen  mutant  among  the  first  generation 
hybrid  progeny  of  O.  laniarckiana  X  biennis  discussed  in  our  earher  account 
(Mutants  and  Hybrids,  p.  18).  The  occurrence  of  this  mutant  along  with 
several  individuals  of  0.  lamarckiana  was  there  attributed  to  an  accidental 
self-fertilization  of  the  pistil  parent,  but  there  is  no  sufficient  reason  perhaps 
for  ruling  out  the  occurrence  of  monolepsis  producing  Millardet  hvbrids  in 
which  the  maternal  characters  alone  appear.  Whether  this  specimen  of  O. 
rubrinervis  was  the  product  of  self-fertilization  or  of  monolepsis  has  no  especial 
significance  for  this  study,  since,  whatever  may  have  been  its  origin,  it  was  a 
true  O.  rubrinervis,  though  having  no  rubrinervis  individuals  in  its  ancestry, 
at  least  for  many  generations. 

The  pollination  of  this  specimen  was  left  entirely  uncontrolled,  and  as  it 
grew  in  close  contiguity  with  other  species  of  Oenothera,  where  insects  were 
busily  engaged  carrying  from  flower  to  flower  the  pollen  of  a  dozen  or  more 
different  species,  the  progeny  might  have  been  expected  to  show  a  confusion 
of  hybrid  forms  of  unknown  affinities.  It  was  a  matter  of  some  surprise, 
therefore,  that  only  i  specimen  in  the  88  appeared  to  be  a  hybrid.  The  char- 
acters of  this  obvious  hybrid  indicated  that  it  was  probably  produced  by  pollen 
from  one  of  the  small-flowered  species,  and  it  was  considered  by  the  senior 
author  to  be  nearlv  if  not  quite  identical  with  his  0.  lamarckiana  X  biennis, 
No.  2.32,  though  no  definite  analysis  of  its  characters  was  made.  The  other 
87  individuals  consisted  of  80  0.  rubrinervis  and  7  which  were  looked  upon  as 
typical  O.  lamarckiana,  although,  as  will  be  seen  later,  the  results  of  the  statis- 
tical studies  present  some  interesting  exceptional  features. 

(rf)  Twenty-seven  individuals  of  O.  rubrinervis  were  grown  from  guarded 
and  self-fertilized  seed  of  one  which  arose  as  a  mutant  at  Amsterdam  in  1895 
in  a  pure  0.  lamarckiana  pedigree  guarded  for  six  generations. 

(e)  Ninety-eight  specimens  of  Oenothera  gigas  were  grown  from  pure-bred 
seed  received  fromProf.  DeVries,  the  pedigree  being  known  and  fully  guarded 
for  five  generations  from  the  time  it  appeared  as  a  mutant  in  1895.  Unfortu- 
nately onlv  4  of  these  came  to  bloom  the  first  year,  so  that  the  studies  here 
given  on  Oenothera  gigas  must  be  considered  wholly  inadequate. 

(/)  Another  lot  of  plants,  44  in  number,  were  the  offspring  of  Oenothera 
lata  which  had  grown  surrounded  by  O.  gigas  in  an  isolated  portion  of  the 
Botanical  Garden  of  Amsterdam.  On  the  side  of  the  pistil-parent  the  pedigree 
of  this  lata  had  been  continued  for  five  generations  from  the  time  it  arose  as 
a  mutant,  and  in  most  of  these  generations  0.  lamarckiana  was  the  pollen  par- 
ent. As  0.  lata  is  nearly  or  quite  incapable  of  self-fertihzation,  owing  to  its 
lack  of  viable  pollen,  these  plants  must  have  been  hybrids  between  0.  lata 
and  O.  gigas,  and  as  they  were  wholly  unselected  they  formed  a  motley  group. 
Among  them  Oenothera  lata  was  the  most  frequent,  17  in  44  belonging  to  this 


20     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF    THE    OENOTHERAS. 

species;  15  were  0.  lamarckiana,  and  2  were  0.  nanella.  The  remaining  10 
were  not  positively  identified,  though  several  which  were  at  first  thought  to 
be  0.  lata  but  which  were  afterwards  seen  to  differ  from  that  species  in  certain 
bud  characters,  especially  in  the  possession  of  pollen-bearing  anthers,  may 
have  been  true  hybrids  between  0.  lata  and  0.  gigas. 

The  advantages  of  bringing  together  to  one  place  for  use  in  this  line  of 
investigation  material  having  such  different  past  histories  will  be  obvious.  If 
the  various  types  of  structure  which  have  been  separated  and  described  under 
specific  names  are  not  perfectly  natural  centers  of  stability,  or  rather  of  equi- 
libration, but  are  the  product  of  selection,  it  is  inconceivable  that  the  forms 
resulting  from  selection  by  different  persons  for  different  periods  of  time  and  in 
different  environments  should  produce  identical  forms,  and  especially  that 
there  should  be  identity  in  characters  that  were  not  taken  into  account  and 
could  not  well  be  taken  into  account  by  those  making  the  selections. 

Some  differences  are  naturally  to  be  expected  in  the  characters  of  these 
several  lots  of  plants  of  the  same  species  coming  from  different  sources, 
because  of  the  known  behavior  of  purely  fluctuating  characters,  and  the  degree 
of  these  differences  will  be  contemplated  with  some  interest.  On  the  other 
hand,  the  various  sources  of  the  material  will  make  the  comparisons  between 
the  different  species  less  satisfactory  than  might  be  wished. 

A  part  of  the  advantage  gained  in  the  way  of  larger  quantities  of  material 
by  bringing  these  cultures  to  the  Station  for  Experimental  Evolution  is  offset 
by  the  fact  that  differences  in  soil,  climate,  and  methods  of  culture  from  those 
to  which  the  cultures  were  subjected  at  the  New  York  Botanical  Garden  in 
1904  will  make  impossible  the  drawing  of  any  conclusions  this  year  regarding 
the  problem  which  originally  incited  these  investigations,  namely,  the  question 
as  to  the  presence  or  absence  of  regression  in  the  mutants.  As  a  partial 
measure  of  the  difference  in  conditions  at  the  New  York  Botanical  Garden  in 
1904  and  at  the  Station  for  Experimental  Evolution  in  1905,  a  comparison  of 
the  heights  of  the  0.  lamarckiana  used  for  the  studies  in  the  former  year  may 
be  made  with  those  of  the  first  lot  (a)  described  on  page  18.  Both  of  these  lots 
were  raised  as  annuals  and  should  therefore  give  a  reasonably  fair  indication 
of  the  relative  value  of  the  resultant  of  all  the  factors  affecting  the  fluctuating 
characters  of  the  two  lots  of  plants. 

The  height-curves  reduced  to  equal  areas  are  shown  superposed  in  fig.  i, 
and  the  principal  constants  are  as  follows : 


New  York  Botanical  Garden,  1004 

Station  for  Experimental  Evolution,  1905. 


Mean. 


Standard 
deviation. 


Centimeters.  I  CenHmeiers. 
88.68  ±o..S5  !  4-76±o.3q 
86.98  ±0.69    8.37±o.j.9 


Coefficient 
of  variation. 


Per  cent. 
5.37  ±0.44 
Q.62  ±0.57 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      21 


It  will  be  noted  that  there  has  been  a  slight  fall  in  mean  heights,  but  the 
most  noteworthy  thing  in  this  comparison  is  the  great  increase  in  the  varia- 
bility. A  study  of  fig.  I  shows  the  difference  in  variabilit)'  quite  clearly  in 
the  greater  amplitude  and  less  height  of  the  1905  curve.  The  interesting 
fact  becomes  apparent  that  although  the  mean  value  of  stem-heights  in  1905 
was  less  than  that  for  1904,  some  of  the  1905  plants  were  taller  than  the  tallest 
in  1904.  The  explanation  of  these  facts  is  very  simple.  The  garden  at  the 
Station  for  Experimental  Evolution  has  been  under  cultivation  so  short  a 


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66-70       71-75      76-80      81-85     86-90     91-95     96-100    101-105    106-110 

Fig.  1. — Variation  in  height  of  stem  of  Oenothera  lamarckiana.  Broken  curve  represents 
the  condition  at  the  Xew  York  Botanical  Garden  in  1904,  continuous  curve  that  at  the 
Station  for  Experimental  Evolution  in  1905. 

time  that  it  still  lacks  much  of  that  homogeneity  that  is  desirable  for  work  of 
this  nature.  One  corner  of  this  plot  of  Oenothera  lamarckiana  dipped  into  a 
slight  depression  which  received  the  drainage  from  the  roof  of  the  laboratory 
and  was  therefore  kept,  during  a  portion  of  the  growing  season,  too  wet  for 
the  best  development  of  the  plants.  A  few  specimens  on  this  corner  did  not 
make  flower-stems  at  all,  and  it  was  to  be  expected  that  those  nearest  this 
corner  of  the  plot  would  be  somewhat  influenced  by  the  same  unfavorable 
conditions  and  made  to  produce  flower-stems  of  a  shghtly  less  altitude. 

The  stature  of  Oenothera   rubrinervis. — In    1904  a  comparison  was  made 
between  the  statures  of  Oenothera  nanella  and  0.  lamarckiana,  and  it  was  a 


22     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF    THE    OENOTHERAS. 


part  of  the  plan  for  the  present  study  that  this  comparison  should  be  contin- 
ued, but  unfortunately  the  seed  of  O.  nanella  gave  only  3  germinations  and 
the  continuation  of  this  comparison  must  be  postponed  until  another  season, 
but  meanwhile  it  will  be  particularly  interesting  to  make  another  comparison 
of  heights  which  will  present  something  of  a  contrast  with  that  between  O. 
nanella  and  its  parent.  It  will  be  recalled  that  in  that  comparison  the  co- 
existence of  two  striking  features  was  taken  to  be  more  than  a  mere  coinci- 
dence, namelv,  the  great  departure  of  the  mean  value  in  0.  nanella  from  the 


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65-70       71-75     76-80       81-85      86-90       91-95      96-100   101-105     106-110      III-II5      116-120 


Fig.   2. — Comparison  of  the  variability  of  stem-heights  of  Oenothera  lamarckiana  (broken  curve) 
and  O.  rubrinervis,  as  grown  at  the  Station  for  Experimental  Evolution  in  1905. 

mean  of  its  parent,  and  a  very  great  increase  in  the  coefficient  of  variability 
in  the  heights  of  the  mutant.  In  comparing  the  heights  of  0.  rubrinervis  with 
those  of  O.  lamarckiana  there  is  seen  a  very  much  less  marked  departure  from 
the  parental  condition  in  the  mean  values,  and  a  correspondingly  slight 
increase  in  the  coefficient  of  variability.  This  comparison  is  shown  graphic- 
ally in  fig.  2  and  is  expressed  numerically  as  follows: 

Variation  in  heights,  1905. 


Mean. 

Standard 
deviation. 

Coefficient 
of  variation. 

Oenothera  lamarckiana 

Oenothera  rubrinervis 

Centimeters. 
86.98  ±0.69 
92. 18  ±0.99 

Centimeters. 
8.37  ±0.49 
9.86  ±0.  70 

Per  cent. 

9.62  ±0.57 

10.  70  ±0.78 

This  may  still  be  merely  a  coincidence,  but  two  such  coincidences  are  very 
much  less  likely  to  occur  than  one,  and  the  attempt  will  be  made  in  the  study 
of  other  characters  to  trace  the  suggested  parallel  between  the  departure  of 
the  mean  from  the  parental  condition  and  the  increase  in  the  value  of  the 
coefficient  of  variabilitv. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS.      23 

The  branching  habit  of  Oenothera  rubrinervis. — In  discussing  the  branching 
of  Oenothera  rubrinervis  in  1904  the  statement  was  made  that  "such  a  char- 
acter as  this  is  too  largely  influenced  by  the  individual  physiological  vigor  to 
be  of  any  value  as  a  diagnostic  character, "  and  this  statement  receives  striking 
confirmation  in  the  conditions  observed  in  1905.  The  specimens  of  0.  rubri- 
nervis grown  at  the  Station  for  Experimental  Evolution  in  the  latter  year  were 
totally  different  in  their  branching  from  those  studied  the  year  before  at  the 


t-.  t/     cd  o>^     o    oj      34-36     37-39    40-42    43-45   46-48 


Fig.   3. — Comparison  of  the  number  of  branches  of  Oenothera  lamarckiana  (shaded  with  lines  rising 
toward  the  right)  and  O.  rubrinervis,  a.s  grown  at  the  Station  for  Experimental  Evolution  in  1905. 

New  York  Botanical  Garden.  There  was  in  1905  almost  a  complete  sup- 
pression of  the  large  basal  branches  which  were  such  a  marked  feature  of  the 
specimens  earlier  described.  Instead  of  finding  Oenothera  lamarckiana  and 
0.  rubrinervis  entirely  distinct  and  separated  by  a  wide  gap  in  respect  to 
number  of  lateral  branches,  the  curves  in  figure  3  show  how  greatly  they 
overlap.     Their  numerical  relations  may  be  studied  in  the  following  table : 


Range. 

Mean. 

Standard 
deviation. 

Coefficient  of 
variation. 

Number  of   the  lateral 

branches : 

Per  cent. 

Oenothera  lamarckiana .  . 

18  to  39 

26.20  ±  o.2>?, 

3-99  ±  0.33 

15-23  ±  0  91 

Oenothera  rubrinervis  .  .  . 

19  to  47 

30  18  ±  0.72 

6.12  ±  0.51 

20.29  ±  1 .76 

Total    branch    length    in 

decimeters: 

Oenothera  lamarckiana .  . 

40  to  150 

91.40  ±  I  49 

17  97  ±  1-05 

19.66  it  1 .20 

Oenothera  rubrinervis  .  .  . 

4T  to  174 

87.22  ±  2  36 

23.46  ±1.67 

26.89  ±  2-05 

24     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 

With  respect  to  the  number  of  branches,  the  figures  in  this  table  do  not 
fully  represent  the  tendency  in  0.  rubrinervis  to  develop  a  large  proportion  of 
its  axillary  buds  into  branches,  as  many  of  these  buds  had  started  into  growth, 
but  for  some  reason  had  been  arrested  before  they  had  attained  a  length  of 
I  cm.,  under  which  length  none  were  counted.  Comparing  these  results  with 
those  of  1904  it  is  found  that  with  respect  to  vanahility  Oenothera  lamarck- 
iana  presents  essentially  the  same  condition  in  the  two  years,  though  the 
values  of  both  number  and  length  of  branches  has  increased  in  the  latter  year. 
In  Oenothera  rubrinervis,  on  the  other  hand,  although  there  has  been  in  1905 
a  great  reduction  both  in  number  and  length  of  branches  and  the  coefficients 
of  variability  are  considerably  changed,  they  are  still  much  higher  than  in  the 
parent  species.  This  will  be  best  appreciated  by  a  comparison  of  the  results 
for  the  two  years  as  shown  in  the  following  table : 


Variation  in — • 

Years. 

Range. 

Mean . 

Standard 
deviation. 

Coefficient 
of  variation. 

Per  cent. 

Number  of  branches  of  ( 

1904 

10  to    27 

20.  27  ±    0.49 

324  ±  0.35 

15.70  lb  1.70 

0.  lamarckiana.          { 

1905 

18  to    39 

26.20  ±    0.33 

.3 

99  ±  0.33 

15-23  ±  0.91 

Number  of  branches  of  < 

1904 

34  to    63 

42.35  ±     1.04 

6 

34  ±  0.73 

1500  ±  1.70 

0.  rubrinervis.              ( 

1905. 

19  to    47 

30. 18  ±    0.  72 

6 

12  ±  0.51 

20.29  it  1 .76 

Branch-length     of    0.  J 

1904 

36  to    85 

66.8    ±     2.0 

13 

5    ±  1.4 

20. 2     ±2.2 

lamarckiana.                ( 

1905 

40  to  150 

914    ±     1-5 

18 

0    ±1.1 

20.0    ±1.2 

Branch-length     of    0.  _< 

1904 

78  to  300 

181. 9    ±  13.0 

79 

5     ±  9  2 

43-7     ±5-1 

rubrinervis.                   ( 

1905 

41  to  174 

87.2     ±     2.4 

23 

5     ±  1.7 

26.9    ±2.0 

The  greater  variability  of  the  mutants  was  taken  in  1904  to  indicate  that 
these  forms  might  not  yet  have  acquired,  perhaps,  that  fixity  of  habit  exhib- 
ited by  an  older  species,  and  the  fact  that  Oenothera  rubrinervis  underwent 
greater  change  in  its  mode  of  branching  on  being  cultivated  in  a  new  environ- 
ment than  was  exhibited  by  0.  lamarckiana  when  subjected  to  the  same 
change  would  accord  well  with  this  suggestion. 

The  increase  in  the  number  and  length  of  the  branches  in  0.  lamarckiana 
and  the  considerable  decrease  in  those  of  O.  rubrinervis  must  not  be  inter- 
preted, however,  as  in  any  sense  an  approach  to  actual  identity  in  these  two 
species,  in  regard  to  branching  habit,  for  in  the  latter  year  they  were  even 
more  distinct  in  general  type  of  branching  than  before.  Oenothera  rubri- 
nervis had  simply  lost  its  long  basal  branches,  while  0.  lamarckiana  had  retained 
them. 


THE  SIZE  AND  SHAPE  OF  LEAVES  OF  OENOTHERA  RUBRINERVIS. 

Some  of  the  difficulties  attending  the  statistical  study  of  leaves  were  pointed 
out  in  our  earlier  paper,  and  the  complete  suppression  of  the  basal  branches  of 
O.  rubrinervis  in  1905  has  made  the  continuation  of  the  comparison  between 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      25 


the  leaves  of  that  species  and  those  of  its 
parent  of  Httle  value  except  as  a  demonstra- 
tion of  the  difficulties  which  were  to  be 
overcome.  It  seemed  fair  in  the  former  year 
to  consider  the  basal  branches  of  these  two 
species  homologous,  and  leaves  for  the  study  95 
in  1904,  being  taken  from  about  the  middle 
of  those  branches,  were  consequently  as 
nearly  comparable  as  possible.  In  1905  the 
leaves  of  0.  lamarckiana  were  again  taken 
from  the  middle  of  the  basal  branches  and 
are  thus  comparable  with  those  of  1904, 
but  in  0.  rubrinervis  the  leaves  were  taken 
from  the  middle  of  the  lowest  branches  pres- 
ent, and  as  these  were  not  quite  basal,  the 
leaves  were  not  homologous  with  those  of 
O.  lamarckiana.  The  first  striking  result 
of  this  lack  of  homology  is  seen  in  fig.  4, 
where  the  variation  in  leaf-lengths  is  com- 
pared. The  corresponding  curves  for  1904 
showed  almost  exact  identity  in  the  length 
of  the  leaves,  while  in  this  figure  it  is  seen 
that  the  length  of  most  of  the  O.  rubri- 
nervis leaves  corresponded  in  1905  with  the 
lower  part  of  the  range  of  0.  lamarckiana. 
When  figure  5,  representing  the  variation  in 
leaf-width,  is  compared  with  the  corres- 
ponding figure  for  1904  it  is  seen  that  with 
respect  to  this  character  the  lack  of  homol- 
ogy has  had  but  little  modifying  effect, 
and  there  is  close  resemblance  between  the 
sets  of  curves  representing  the  variation  in 
leaf-width  for  the  two  years,  though  both 
species  show  a  slight  increase  in  the  coef- 
ficient of  variability.  The  considerable 
difference  in  respect  to  leaf-length,  taken 
in  conjunction  with  essential  identity  in 
leaf- width  in  1905,  has  the  effect  to  change 
the  ratios  between  length  and  width  in 
about  the  same  degree  as  the  change  in 
length,  but  in  the  opposite  direction.  Thus 
the  change  in  length  was  away  from  identity, 


IIO 


105 


100 


90 


85 


80 


75 


70 


65 


60 


55 


50 


45 


35 


30 


25 


20 


15 


10 


( 1 

1 

0 

<1 

?f\     0 

''\\°  '"' 

'1      ''    ! 
'    ,      0   ', 

0 

1     "         ( 
f    ° 

>  ', 

6 

1 

II 

0 

T    ' 

1 

0 

\ 

c 

) 

i 

J4 

I 

A 

4-6 
50 


71 
75 


96 
100 


121 
125 


14-6 
150 


Fig.  4. — Comparison  of  leaf -length  of  Oeno- 
thera lamarckiana  (broken  curve)  and  O. 
rubrinervis. 


26     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS. 


while  that  in  the  leaf 
ratios  was  toward  iden- 
tity, as  will  be  readily 
seen  on  referring  to  fig. 
6, which  shows  a  much 
more  complete  over- 
lapping than  do  the 
corresponding  curves 
for  1904.  In  other 
words,  since  this  ratio 
was  taken  as  a  measure 
of  leaf-form,  the  form 
of  the  leaves  of  O. 
rubrinervis  in  the  latter 
year  was  but  slightly 
different  from  that  of 
the  parent  species,  ex- 
cept in  those  minor 
details  which  were 
pointed  out  in  the 
earlier  study  as  pre- 
venting the  ratio  be- 
tween length  and 
width  from  being  a 
correct  measure  of 
form.  The  lack  of 
homology  in  the  leaves 
measured  in  the  two 
species  takes  away  all 
significance  of  these 
facts  in  their  bearing 
on  the  question  of  the 
regression  of  0.  rubrinervis.  The  constants  of  the  several  curves  representing 
the  variation  in  leaf  measurements  may  be  compared  in  the  following  table : 


Fig.   5.- 


-Comparison  of  leaf -width  of  Oenothera  lamarckiana  (broken  curve) 
and  O.  rubrinervis. 


Variation  in  length  of  leaf: 

Oenothera  lamarckiana 

Oenothera  rubrinervis 

Variation  in  width  of  leaf: 

Oenothera  lamarckiana 

Oenothera  rubrinervis 

Variation  in  the  ratio  between  width  and 
length : 

Oenothera  lamarckiana 

Oenothera  rubrinervis 


Mean. 

Millimeters. 
88.91  ±0.35 
70.96  ±0.24 

36.56±o. 
27.3o±o. 

091 
079 

Per  cent 
42.27  ±0 
39-34  ±0 

! 
10  t 
13 

Standard 
deviation. 


Millimeters. 

18.41  ±0.25 

11.42  ±0.17 

4.  7q6  ±o.o6j 
3.687  ±0.056 

Per  cent. 
5.411  ±0.072 
5.888  ±o.o8q 


Coefficient 
of  variation. 

Per  cent. 
20.71  ±0.29 
16.  io±o.24 

13.  i2±o.  18 

13    50±0.2I 


1 2 . 80  ±  o. 1 7 
i4-97±o.23 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      27 


The  original  measurements  upon  which  the  above  figures  and  conclusions 
are  based  are  presented  in  the  form  of  correlation  tables  in  figs.  7  and  8.  A 
comparison  of  the  tables  with  the  corresponding  figures  for  1904  shows  that 
those  representing  0.  lamarckiana  are  very  much  alike  in  the  two  years,  the 
degree  of  correlation  being  even  higher  in  the  latter  year  than  before,  due  in 
part  possibly  to  the  slightly  greater  heterogeneity  already  pointed  out  in  the 
cultures  at  the  Station  for  Experimental  Evolution.  On  the  other  hand,  in 
Oenothera  rubrinervis  the  corresponding  tables  for  the  two  years  are  entirely 
50r 


Fig.   6. — Comparison  of  the  ratio  between  leaf-length  and  leaf-width  in  Oenothera 
lamarckiana  (broken  curve)  and  O.  rtibrinerzis. 

different,  as  would  be  expected  from  what  has  already  been  said.     In  this 
species  the  coefficient  of  correlation  has  been  considerably  lowered. 

The  coefficients  of  correlation  for  the  two  species  for  the  two  years  may 
be  compared  in  the  following  table : 


1904. 

1905. 

Oenothera  lamarckiana 

0.7916  ±  0.0090 
0.6604  ±  0.0119 

0.8330  ±  0.0058 
0.4452  ±  0.0171 

Oenothera  rubrinervis 

28     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

Variation  in  bud  characters. — It  is  a  well-known  fact  that  floral  characters 
fluctuate  less  in  response  to  change  of  environment  than  vegetative  characters, 
though  I  know  of  no  exact  studies  showing  to  what  extent  this  is  true.  The 
comparatively  larger  flowers  of  alpine  regions  find  an  explanation  on  this 
basis  since  the  alpine  plants  have  generally  migrated  from  the  lowlands. 
I  have  seen  a  specimen  of  Lobelia  syphilitica  less  than  8  cm.  tall  and  dwarfed 


Xo 

-9 

-8-7 

-6 

—  5 

-4 

-3 

-o 

-i 

0 

1 

2 

3 

4 

5 

G     7 

8 

9 

10 

11 

12 

13 

10 

15   50 

55 

60 

65 

70 

75 

80 

85 

90 

95 

100 

105 

110115!l2o'l25 

130'l35 

140 

145 

150 

45 

50  55 

60 

65 

70 

75 

80 

85 

90 

95 

100105 

110 

115120125 

130135IU0 

145 

150 

155 

Xi 

- 

-13 

23 

1 

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-12 

24 

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3 

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25 

3 

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26 

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8 

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-8 

28 

1 

10 

10 

7 

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31 

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2'J 

3 

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7 

9 

4 

1 

31 

-G 

30 

2 

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10 

9 

9 

1 

13 

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31 

1 

/ 

13 

9 

5 

3 

3 

3 

11 

-4 

32 

5 

12 

14 

27 

G 

8 

6 

3 

81 

-3 

33 

2 

10 

12 

21 

13 

19 

12 

C 

1 

2 

98 

—2 

34 

0 

12 

20 

22 

19 

10 

G 

4 

5 

1 

105 

-1 

35 

1 

4 

3 

IG 

10 

27 

18 

13 

10 

4 

2 

108 

0 

36 

G 

3 

5 

8 

14 

10 

181  a 

5 

1 

79 

1 

37 

G 

11 

7 

14 

21 

20 

10 

8 

G 

2 

105 

2 

38 

2 

3 

G 

9 

12 

26 

11 

21 

4 

4 

6 

1 

105 

3 

39 

2 

3 

1 

3 

0 

8 

iG 

18 

10 

3 

2 

3 

75 

4 

40 

1 

2 

2 

1 

5 

17 

15 

15 

14 

5 

3 

5 

85 

5 

41 

2 

2 

4 

4 

7 

8 

12 

8 

3 

3 

53 

6 

42 

1 

1 

1 

3 

10 

11 

11 

7 

11 

1 

2 

1 

CO 

7 

43 

1 

2 

4 

4 

7 

4 

11 

6 

2 

1 

1 

43 

8 

44 

1 

3 

9 

9 

4 

1 

27 

9 

45 

2 

4 

10 

5 

4 

1 

1 

2 

29 

10 

46 

1 

^ 

3 

3 

2 

9 

1 

16 

11 

47 

1 

O 

3 

O 

2 

1 

1 

12 

12 

48 

1 

1 

1 

1 

1 

13 

49 

3 

1 

1 

5 

14 

50 

o 

1 

1 

o 

1 

7 

15 

51 

1 

1 

2 

1 

2     7 

33 

75 

91 

111 

91 

121 

110129 

98 

103 

SI 

63 

53 

35 

12 

8 

6 

2 

0 

2 

1270 

Fig.   7. — Correlation  between  width  (subject)  andlength  oi\eaV\nOcnothi:ralama)ckiana. 
Measurements  in  millimeters.      Coefficient  of  correlation,  0.8330  i  0.0058. 

specimens  of  numerous  other  species  bearing  flowers  almost  as  large  as  is 
normal  for  the  species. 

Having  sufficiently  demonstrated  the  unsatisfactory  character  of  height, 
branching,  and  leaf-form  for  statistical  comparisons  because  of  their  great  sen- 
sitiveness to  even  shght  changes  in  the  environment,  I  have  decided  that  in  the 
buds  is  to  be  found  probably  the  most  satisfactory  material  for  a  continuation 


PLATE  8. 


Huds  of  ()enotlier;i  himarckiana  and  four  of  its  mutants,  showing:  characteristic  differences  and 
variability,  ist  row  at  top,  Oenothera  lamarckiana;  2nd  row,  Oenothera  ruhrinervis;  3rd 
row,  Oenothera  lata ;  4th  row,  left,  Oenothera  gigas,  right,  Oenothera  nanella. 


5i 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      29 

of  these  studies.  The  oenotheraceous  bud  offers  distinctive  characters  for 
the  separation  of  the  several  species  (plate  8).  It  has,  as  is  well  known,  three 
distinct  regions,  the  ovary,  the  hypanthium,  and  the  cone — which  are  in  most 
cases  marked  off  from  each  other  in  such  a  way  as  to  make  their  accurate 
measurement  easy.  As  proof  of  this  fact  it  is  only  necessary  to  say  that  fre- 
quent repetitions 
of  the  measure- 
ments showed  only 
a  variation  of  one- 
tenth  of  a  milli- 
meter in  the  thick- 
ness of  ovary  and 
hypanthium,  and 
from  o.2too.3mm. 
in  thickness  of  the 
cone  and  lengths  of 
the  several  regions, 
the  maximum  error 
in  the  smallest 
me  asurements 
taken  being  less 
than  4  per  cent  of 
the  magnitude 
measured,  and  in 
the  larger  measure- 
ments,  such  as 
length  of  hypanthi- 
um and  cone,  less 
than  1  per  cent. 

In  deciding  to 
make  statistical 
studies  on  buds  a 
careful  preliminary 
survey  was  made 
to  determine  what 
precautions  would 
need   to  be  taken 

in  order  to  free  the  work  from  error.  The  several  points  that  seemed  to  need 
consideration  were :  (a)  A  probable  periodicity  in  the  size  of  the  buds,  so  that 
those  in  the  early  part  of  the  season  are  doubtless  a  little  larger  than  those 
produced  near  the  end  of  the  season,  but  as  the  flowering  season  is  very  long  this 
periodicity  is  certainly  slight  and  would  probably  have  no  appreciable  influence 


1    1 

X„ 

-4 

-3 

-2  -1 

0 

1 

2 

3 

4 

5     6 

7     8 

9 

10 

u 

12 

13 

14  15 

i5 

50 

55 

GO 

65 

70 

75 

80 

85 

90  95 

100|105 

110 

115 

120 

125 

130 

135 

HO 

50 

00 

60|05 

70 

75 

80 

85 

00 

95  100 

105 

110 

115 

120 

125 

130 

135 

140 

145 

Xi 

-11 

16 

1 

1 

-10 

n 

0 

-9 

18 

1 

1 

1 

1 

1 

-8 

19 

1 

1 

1 

1 

1 

1 

6 

-7 

20 

4 

1 

4 

4 

1 

2 

le 

-G  21 

|3 

7 

9 

0 

0 

2X 

-5,22 

2  1  3  !  13 

12 

7 

5 

1 

43 

-4|23 
-3j24 

15  [14 

16 

9 

5 

1 

4 

I 

55 

1     5  1 12, 12 i 13 

5 

0 

4 

2 

2 

59 

-2125 

'      !  3 

23  20 

33 

17 

13 

6 

1 

1 

1 

118 

-I 

26 

i      1  -1 

10  29 

19 

15 

5 

8 

5 

95 

0 

27 

1  j  4  !  9  '21' 31 

18 

14 

7 

G 

4 

0 

0 

119 

1 

28 

1      !  S  .  18 , 2'J 

32 

10 

10;  G 

1 

5 

1 

120 

2 

29 

I  1 

10  [16 

OO 

5 

5 

3 

2 

3 

07 

13 

30 

1  '  1  '  2 

9 

18 

13 

12 

7 

6 

4 

1 

2 

1 

2 

79 

I4 

31 

1  1 

4 

7 

III  12 

11 

6 

3 

1 

1 

1 

1 

59 

5 

32 

3 

1 

G 

12 

S 

3 

4 

0 

3 

39 

G 

33 

I 

1 

1 

7 

7 

13 

7 

3 

1 

1 

2 

1 

1 

4G 

7 

34 

2 

4 

3 

4 

4 

1 

IS 

8 

35 

2 

1 

3 

2 

1 

1 

3 

13 

9 

36 

2 

1 

2 

1 

6 

10 

.37 

1 

1 

1 

1 

i 

11 

38 

1 

1 

2 

12' 39 

2 

2 

13  40 

0 

h'ji 

0 

15 

42 

0 

16 

43 

0 

17 

44 

0 

18 

45 

0 

19 

46 

0 

20 

47 

1 

1 

7 

3t 

lOfl 

172 

209il7l 

1U2 

S3 

47 

21 

17 

12 

2 

3 

0 

1 

1 

0 

0 

1 

995 

1 

P'lG.  8. — Correlation  between  width  (subject  )and  length  of  leaf  in  Oenolliera 
rubrinenis.  Measurements  in  millimeters.  Coefficient  of  correlation, 
0.4452±0.0171. 


30     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


during  the  greater  part  of  the  season  if  the  earUest  and  latest  flowers  are 
omitted,  (b)  A  variation  in  the  size  of  the  buds  in  response  to  different  con- 
ditions of  the  weather.  While  this  variation  is  not  often  great  enough  to  be 
noticeable,  there  were  times  at  which  the  buds  seemed  distinctly  larger  after 
several  days  of  warm,  humid  weather.  Both  of  these  difficulties  would  be 
rendered  comparatively  harmless  by  a  collection  of  a  small  number  of  buds 
each  day  from  each  of  the  lots  of  plants  studied,  so  that  all  were  being  subjected 
to  the  same  conditions.  There  are  still  some  elements  of  error,  however, 
which  this  method  does  not  fully  meet,  such,  for  instance,  as  the  existence  of  a 
different  period  in  the  different  species,  or  a  different  kind  or  degree  of  reaction 
to  changes  in  the  weather.     Without  an  extensive  study  of  the  periodicity  and 

response  in  the  several 
species  to  changed  cli- 
matic conditions  it  can 
not  be  known  just  how 
serious  errors  these 
factors  introduce,  but 
it  is  believed  that  they 
are  slight,  (c)  There 
is  always  present  a 
complete  series  of 
buds,  beginning  with 
those  ready  to  bloom 
and  ending  with  those 
just  beginning  to  de- 
velop, and  the  develop- 
ment is  so  rapid  that 
buds  collected  at  one 
hour  of  the  day  are 
not  the  equivalent  of  those  collected  at  any  other  hour.  It  was  found  on 
examination  that  the  buds  which  would  bloom  the  following  day  could  be 
readily  distinguished  by  their  size  and  color.  Rarely  there  would  be  a  doubt 
about  some  particular  bud,  and  such  were  omitted  from  the  study.  To 
avoid  the  collection  of  buds  at  different  stages  of  development  they  were 
collected  from  all  the  lots  at  5  p.  m.  and  measured  as  soon  thereafter  as  pos- 
sible. This  makes  all  the  buds  used  in  these  studies  as  nearly  comparable  as 
could  well  be  secured,  though  it  is  realized  that  a  source  of  slight  error  is  to  be 
found  in  the  fact  that  on  some  days  the  development  of  the  buds  is  more 
rapid  than  on  others,  so  that  they  are  not  always  quite  equivalent  at  the  time 
of  collection,  when  this  is  determined  by  the  clock  without  regard  to  the 
weather. 


Xo 

-11 

-10 

-9 

-8 

-7 

-6 

-5 

-4 

-3 

_o 

-1 

0 

1 

2 

3 

4 

5 

G 

7 

o» 

•23 

24 

■25 

26 

•1- 

28 

29 

30 

31 

32 

33 

34 

3o 

3i; 

37 

38 

39 

40 

Xi 

-6 

60 

65 

1 

1 

-5 

65 

70 

0 

-4 

70 

75 

1 

1 

1 

1 

1 

5 

-3 

75 

80 

1 

2 

I 

1 

5 

_2 

80 

S5 

1 

1 

5 

2 

1 

3 

2 

1 

4 

1 

0 

1 

1 

25 

-1 

85 

00 

1 

1 

1 

3 

1 

4 

1 

0 

5 

3 

1 

23 

0 

90 

95 

1 

1 

0 

1 

4 

2 

1 

5 

3 

4 

0 

2 

0 

30 

1 

05 

IOC 

2 

1 

3 

0 

0 

5 

1 

G 

4 

4 

5 

5 

1 

1 

42 

2 
3 

100 

1C5 

1C6 
110 

1 

1 

1 

4 

3 

2 

3 

2 

17 

2 

1 

4 

1 

2 

10 

i 

110 

115 

1 

1 

5 

115 

120 

1 

1 

2 

1 

1 

0 

4 

S 

9 

5 

8 

14 

12 

5 

IS 

20 

10 

13 

17 

8 

3 

iGd 

Fig.  9. — Correlation  in  the  buds  oi  OenotheralamarckianaiTomHilveT- 
sum.  Length  of  ovary  subject,  length  of  hypanthium  relative. 
Coefficient   of   correlation,  0.3872  zb  0.0453. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      3 1 

This  work  was  not  begun  until  late  in  the  season,  the  first  collection  of  buds 
for  this  study  having  been  made  on  September  1 5  and  the  last  on  October  30, 
and  this  will  need  to  be  taken  into  account  in  making  future  comparisons, 
though  it  seems  likely  that  the  periodicity  is  too  slight  to  make  this  a  modify- 
ing factor  of  any  considerable  moment. 

The  number  of  buds  from  each  lot  of  specimens,  except  one,  was  from  140 
to  161.  In  Oenothera  gigas  only  91  buds  were  secured  before  frost  cut  short 
the  investigation.  The  lateness  of  the  time  at  which  the  work  was  begun 
prevented  the  measurement  of  a  larger  number  of  buds  of  each  lot,  though  it  is 
realized  that  a  much  larger  number  would  be  desirable.  On  this  account  it 
is  not  thought  necessary  to  present  variation  curves  of  each  lot  of  measure- 
ments. Instead,  the  constants  have  been  calculated  and  are  presented  in  the 
form  of  a  table : 


Length  of  ovary. 

Length  of  hypanthium. 

Mean. 

Standard 

Coefficient 

Mean. 

vStandard 

Coefficient 

deviation. 

of  variation. 

deviation. 

of  variation. 

Deci-millimeters. 

Deci-  m  illimete  rs 

Per  cent. 

Milliineters. 

Millimeters. 

Per  cent. 

a 

95  67    ±0  50 

9-33   ±0.35 

9-75±o.39 

3306    ±0.23 

4.20    ±0.16 

I2.7I±0.49 

b 

91    76    ±0.47 

S.73    ±0.33 

9.52±o.36 

33  07   ±0.22 

4 

18    ±0.16 

i2.63±o.48 

c'* 

92.29    ±0    34 

6  40   ±0.24 

6  94±o.26 

29.10    ±0.22 

4 

14   ±0.16 

i4.23±o.55 

c 

10756    ±0.53 

9.89   ±0.37 

9-20±o.35 

30.37    ±0.21 

3 

88    ±0.15 

i2.77±o.49 

d 

Ti8.8i    ±0.47 

8.40   ±0.33 

707±o.  28 

30.33    ±0.18 

3 

24    ±0.13 

io.69±o.43 

e 

102.12    ±0.64 

9.01    ±0.-45 

8.83±o.44 

25-18    ±0.37 

5 

29    ±0.26 

2 1 . 00  ± 1 . 09 

f 

83.42    ±0.44 

7.77    ±0.31 

9-3i±o.38 

26.96    ±0.32 

5 

61    ±0.23 

2o.8o±o.87 

Length  of  cone. 

Thickness  of  ovary. 

Millimeters. 

Millimeters. 

Deci-millimeters. 

Deci-millimeters 

a 

39-47    ±0.19 

3-47    ±0.13 

8.79±o.34 

32-37     ±0.10 

I.  932  ±0.073 

5-97±o.23| 

b 

39 

47    ±0.18 

^ 

45    ±0.13 

8.74±o.33 

33.51     ±0.11 

2.o78±o.078 

6 

20±0.24 

c' 

37 

80   ±0.14 

2 

65    ±0.10 

7  .01  ±0.26 

31  .261  ±0.091 

1 .711  ±0.064 

5 

47±o.2i 

c 

35 

52    ±0   17 

3 

29    ±0.15 

9.26±o.35 

33-76    ±0.14 

2  .540  ±0.096 

7 

52±0.28 

d 

36 

35    ±0.13 

2 

277  ±0.090 

6.26±o.25 

36.48     ±O.TI 

2.041  ±0.081 

5 

59  ±0  22 

e 

39 

51    ±0.36 

5 

29    ±0.26 

i2.75±o.65 

44.46    ±0.31 

3.69  ±0.22 

8 

3o±o.49 

t 

30 

27    ±0.24 

4 

27    ±0.17 

i409±o.58 

38.24    ±0.11 

i.938±o.o78 

5.o7±o  20 

— 

Thickness  of  hypanthium. 

Thickness  of  cone. 

Deci-millimeters.   \ Deci-millimeters 

Deci-millimeters. 

Deci-millimeters 

a 

25   291  ±0.068  r  .261  ±0.048 

4-99±o   19 

84.59    ±0   39 

7.21     ±0.27 

8.53±o.33 

b 

26.o69±o.o7o 

I 

.32o±o.o5o 

5.o6±o.  19 

81 

81    ±0.32 

6.00    ±0.23 

7 

33±o.28 

c' 

25-453±o.o74 

I 

389±o.o52 

5-46±o.2i 

87 

60    ±0.35 

6.63    ±0.25 

/ 

56±o.29 

c 

28.46   ±0.  10 

I 

940  ±0.073 

6.82±o.26 

93 

26    ±0  43 

8.01     ±0.30 

8 

59±o.33 

d 

30-356±o.o70 

I 

259 ±0  050 

4-  i5±o.i6 

93 

47    ±0.29 

5.17     ±0.20 

5 

53±o.2i 

e 

36.45   ±0.24 

3 

46   ±0.17 

9. 48  ±0.48  109 

76   ±0.69 

9-79    ±0.49 

8 

92±o.45 

f 

29-  i57±o.o8i 

I 

422±o.o57 

4.88±o  20 

96 

41    ±0.47 

8.18    ±0.33 

8.48±o.34 

*cf  ^Oenothera  lamarckiana  from  lot  c. 


32     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

One  of  the  things  of  first  interest  that  will  be  looked  for  in  this  table  is  the 
relation  of  the  fluctuation  found  in  different  lots  of  plants  of  the  same  species. 
Particularly  is  it  interesting  to  compare  the  conditions  of  entry  a  with  entry 
b,  the  first  representing  Oenothera  lamarckiana  self-fertilized  for  sixteen  genera- 
tions, the  second  the  same  species  which  has  never  been  self-fertilized,  except 
as  that  process  may  have  taken  place  by  natural  causes. 

One  represents  i6  generations  of  artificial  selection,  the  other  many  genera- 
tions of  natural  selection,  but  in  neither  case  is  it  probable  that  the  propor- 
tionality in  the  parts  of  the  bud  entered  into  the  selection  in  any  specific  way. 
In  the  mean  values  of  the  several  parts,  these  two  lots  of  plants  are  found  to  be 


1 

Xo 

-13 

-VI 

Ui 

-10 

-9 

-8 

~  i 

-G 

-5 

-4 

-3 

—  o 

-1 

0 

1 

2 

3 

4 

5 

G 

7 

8 

9 

10 

20 

21 

22 

23 

24 

25 

20 

27 

28 

29 

30 

31 

32 

33 

34 

35 

3G 

37 

38 

39 

40 

41 

42 

43 

X, 

-11 

35 

40 

1 

1 

-10 

to 

45 

0 

-y 

45 

50 

0 

-s 

50 

55 

0 

~< 

55 

60 

0 

-fi 

60 

65 

0 

-5 

65 

70 

0 

-4 

70 

75 

0 

-3 

75 

SO 

1 

1 

2 

4 

_o 

SO 

85 

1 

2 

3 

-1 

0 

85 
90 

90 

95 

2 

1 

1 

o 

2 

2 

1 

3 

5 

2 

1 

•n 

1 

1 

1 

O 

2 

4 

1 

3 

4 

G 

4 

o 

5 

1 

1 

42 

1 

95 

lUO 

1 

2 

1 

3 

1 

2 

3 

4 

2 

6 

i 

2 

1 

1 

1 

3S 

o 

100^05 

1 

1 

2 

3 

1 

2 

o 

o 

2 

2 

19 

3 

lOO'llO 

2 

1 

2 

4 

1 

1 

3 

1 

1 

17 

4 

110 

115 

1 

1 

2 

1 

2 

7 

5 

115 

120 

1 

1 

1 

3 

G 

120 

125 

1 

1 

I 

0 

1 

0 

O 

4 

4 

4 

4 

13 

7 

11 

17 

20 

8 

12 

13 

15 

7 

7 

4 

3 

0 

1 

158 

Fig.  10. — Correlation   in   the  buds  of  OenoUiera   lamarckiana,    Xo.  04129.     Length  of 
ovary  subject,  length  of  hypanthium  relative.      Coefficient  of  correlation,  0.4215  ± 


0.0441. 


identical  only  in  regard  to  length  of  hypanthium  and  length  of  cone.  Consid- 
erable differences  are  to  be  seen  in  length  of  ovary  and  thickness  of  cone,  and 
slight  differences  in  thickness  of  ovary  and  of  hypanthium.  In  the  standard 
deviation  and  coefficient  of  variability,  on  the  other  hand,  there  is  essential 
identity  in  every  character  except  in  the  thickness  of  the  cone,  which  was 
significantly  more  variable  in  the  self-fertilized  plants.  There  are  too  many 
possible  sources  of  such  difference  to  make  any  speculation  regarding  its 
significance  profitable  at  this  time. 

When  the  data  in  the  third  entry  (c'),  is  compared  with  those  in  the  first 
and  fourth  entries,  the  very  interesting  fact  develops  that  in  certain  charac- 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE)    OENOTHERAS.      33 


Xo 

-9 

-S 

~  J 

-G 

-5-4 

-3 

—2 

-1 

0 

1 

0 

3 

4 

5 

0 

7 

S 

9 

lu 

U 

■20 

•71 

00 

■23 

24  '25 

■2G 

27 

2S 

29 

30 

31 

32 

33 

34 

35 

3G 

37 

3S 

39 

40 

X, 

-4 

70 

75 

1 

1 

"■J 

75 

80 

0 

—2 

SO 

85 

1 

0 

2 

1 

2 

2 

1 

1 

I 

2 

15 

-1 

Ho 

90 

I 

1 

3 

/ 

G 

5 

3 

4 

3 

i 

1 

5 

1 

3 

47 

0 

uo 

<J5 

1 

1 

1 

2 

4 

- 

4 

1 

1 

3 

3 

0 

3 

1 

0 

1 

40 

1 

95 

100 

3 

3 

2 

4 

•J 

5 

5 

0 

2 

'> 

5 

37 

0 

100 

105 

1 

1 

2 

1 

1 

1 

1 

1 

1 

1 

1 

12 

3 

105 

IIU 

1 

1 

1 

2 

2 

1 

1 

9 

0 

3 

2 

5 

1-2 

8 

15 

16 

15 

8 

15 

lU 

14 

8 

11 

9 

2 

3 

1 

1 

1 

^61 

Fig.  11  .—Correlation  in  the  buds  of  Oenothera  lamarckiana.  No.  04113. 
Length  of  ovary  subject,  length  of  hypanthium  relative.  Coefficient  of 
correlation,  0.4719  ±  0.041.?. 


ters,  as  thickness  of  hypanthium  and  length  and  thickness  of  cone,  the  plants 
here  represented  stand  intermediate  between  those  of  a  and  c,  and  that  the 
mean  length  of  hypanthium  was  even  less  than  in  either,  though  it  should 
have  been  expected 
to  be  as  great  as 
that  of  a ;  in  other 
words,  the  speci- 
mens oiO.  lamarck- 
iana which  sprang 
from  a  rubrinervis 
parent  had  certain 
characters  interme- 
diate between  pure- 
bred 0.  lamarckiana 
and  O.  rubrinervis, 
and  in  length  of 
hypanthium  they 
were  even  less  than 

those  of  the  latter  species.  These  plants  were  considered  typical  Oenothera 
lamarckiana,  not  only  by  myself,  but  by  others  who  are  familiar  with  that 
species.  Here  again,  as  in  the  rest  of  the  data  presented  in  this  table,  the 
general  validity  of  any  conclusion  reached  may  be  questioned  because  of 
the  small  amount 
of  data. 

Comparison  of 
the  entries  c  and 
d,  representing  the 
variation  in  the 
buds  of  two  dif- 
ferent pedigrees  of 
0.  rubrinervis, 
shows  that  only  in 
length  of  hypan- 
thium and  thick- 
ness of  cone  were 
the  two  lots 
essentially  alike 
in  mean  value, 
the  means  of  the 
other  parts  differ- 
ing from  each  other  by  more  than  the  sums  of  the  probable  errors.  In  respect 
to  variability,  entry  c  shows  considerably  greater  coefficients  in  every  set  of 


~ 

Xo 

rll 

-10 

-9 

-8 

—7 

-6 

-5 

-•4 

-3 

_  0 

-1 

0 

1 

2 

3 

4 

5 

G 

7 

8 

9 

10 

•2is 

•26 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

4G 

47 

48 

49 

Xi 

-6 

60 

C5 

1 

1 

-5 

05 

70 

0 

-4 

Hi 

75 

I 

1 

1 

1 

1 

5 

-3 

75 

80 

2 

2 

1 

5 

_o 

80 

85 

1 

2 

4 

3 

3 

1 

3 

4 

2 

0 

25 

-1 

85 

90 

0 

2 

2 

2 

G 

2 

2 

2 

1 

1 

1 

OC 

0 

90 

95 

1 

1 

5 

1 

3 

•■» 

4 

5 

5 

-' 

1 

3J 

1 

95 

100 

1 

2 

'> 

4 

4 

3 

7 

<i 

5 

5 

5 

1 

42 

3 

IOC 

1C5 

1 

2 

2 

3 

0 

1 

3 

1 

1 

1 

17 

1C5 

no 

1 

1 

1 

5 

1 

1 

10 

4 

no 

115 

1 

1 

5 

115 

ICO 

1 

1 

1 

1 

0 

0 

1 

1 

7 

7 

IG 

11 

19 

10 

20 

IG 

13 

12 

11 

7 

2 

0 

1 

0 

lUO 

Fig.  12. — Correlation  in  the  buds  of  Oenothera  lamarckiana  from  Hilversum. 
Length  of  ovary  subject,  length  of  bud-cone  relative.  Coefficient  of  correla- 
tion, 0.6340±  0.0319. 


34     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

measurements  than  entry  d.  The  two  lots  of  plants  here  compared  may  be 
contrasted  with  respect  to  their  origin  and  treatment  as  follows,  and  seme 
one,  several,  or  all  of  these  contrasted  factors  will  probably  satisfactorily 
account  for  the  difference  in  variability: 


c. 

d. 

Seed  from  the  New  York  Botanical  Gar- 
den. 

Of  hybrid  (?)  origin,  0.  lamarckiana  X 
biennis. 

PolHnation  not  guarded. 

Seeds  sown  Feb.  i6,  1905. 

Buds  collected  Sept.  15  to  28,  being  the 
last  half  of  the  season. 

Seed  from  the  Botanical  Garden  of  Am- 
sterdam. 
From  pure-bred  0.  lamarckiana. 

Pollination  guarded,  self-fertilized. 
Seeds  sown  Mar.  11,  1905. 
Buds  collected  Sept.  30  to  Oct.  10,  being 
early  in  the  flowering  season. 

It  remains  to  compare  the  three  mutants  here  represented — 0.  ruhrinervis, 
0.  gigas,  and  0.  lata — ^with  the  parent  species  in  regard  to  mean  values  of 
their  several  characters  and  the  coefficients  of  their  variability.     This  can 

not  be  done  with  entire 
satisfaction  because  of  the 
inconsistencies  in  the  meas- 
urements of  plants  belong- 
ing to  different  pedigrees 
in  the  same  species.  Such 
comparison  would  be  just 
only  when  the  lots  of  mate- 
rial of  all  the  species  had 
received  identical  treat- 
ment, and  this,  as  we  have 
seen,  has  not  been  true  of 
these.  Wherever  several 
lots  of  plants  of  a  single 
species  have  been  used,  the 
average  condition  of  the 
several  lots  has  been  taken 
for  comparison,  and  as  a 
given  amount  of  deviation 
in  a  constant  of  one  value 
is  unfairly  compared  with 
a  given  deviation  from  a 
constant  of  a  different  value,  all  the  deviations  have  been  reduced  to  per  cents 
of  the  equivalent  measurement  in  Oenothera  lamarckiana.  These  values  are 
arranged   in   the   following  table  in   such  a  way  as  to  allow  a  comparison 


~ 

r 

x. 

-7 

-(1 

-5 

-1 

-3 

-2 

-1]  0 

1 

0 

:; 

4 

.5 

6 

7 

8 

9 

32 

33 

31 

35 

30 

37 

38 

3'J 

W 

11 

12 

13 

44 

45 

46 

47 

48 

Xi 

-11 

3^ 

to 

1 

1 

-10 

10 

15 

0 

-9 

Ij 

50 

0 

-h 

5(J 

55 

0 

-7 

o5 

60 

0 

-6 

60 

65 

0 

-5 

65 

70 

0 

— ! 

70 

75 

0 

-3 

75 

80 

1 

1 

1 

1 

4 

0 

80 

85 

1 

1 

I 

3 

-1 

85 

90 

2 

1 

3 

3 

3 

5 

3 

1 

2 

23 

0 

91, 

95 

1 

-1 

1 

10 

6 

9 

A 

2 

2 

2 

1 

42 

1 

95 

100 

I 

1 

1 

3 

5 

5 

5 

7 

3 

4 

0 

1 

38 

.-) 

ICO 

105 

1 

1 

0 

0 

0 

4 

1 

1 

1 

19 

3 

105 

110 

0 

1 

1 

1 

3 

4 

1 

1 

17 

4 

110 

115 

1 

1 

1 

1 

2 

7 

5 

115 

120 

1 

1 

3 

6 

120 

125 

1 

1 

— 

1 

3 

2 

8 

9 

■22 

19 

20 

16 

11 

17 

0 

5 

6 

4 

3 

3 

158 

_J 

Fig.  13. — Correlation  in  the  buds  of  Oenothera  lamarckiana.  No. 
04129.  Length  of  ovary  subject,  length  of  bud-cone  relative. 
Coefficient  of  correlation,  0.5631  ±  0.0366. 


MUTATIONS,    VARIATIONS,    AND   REIyATIONSHIPS   OF   THE    OENOTHERAS.      35 

between  the  deviation  in  the  mean,  dM,  and  that  of  the  coefficient  of  vari- 
abihty,  dV ,  not  only  of  the  characters  studied  in  1905,  but  also  those  of 
the  preceding  year  derived  in  the  same  way. 


Deviation  of  means  and  variabilities,  in  per  cents  of  the  corresponding  values  for  Oenothera 

lamarckiana. 


1904. 


O.  rubrinervis 
Length  of  leaf .  , 

Width  of  leaf 

I-eaf  ratio 

Number  of  branches. 
Total  branch  length . 

O.  nanella,  1904. 
Height 


O.  rubrinervis,  igo$. 

Length  of  leaf 

Width  of  leaf 

Leaf  ratio 

Height 

Number  of  branches. .  . 
Total  branch  length    .  . 

Length  of  ovary 

Length  of  hypanthium. 
Length  of  cone 


dM 

dV 

1 .02 

-   7.21 

20 .  95 

6.91 

24-75 

8.08 

104-59 

-4.46 

172.30 

116.34 

74.28 

492 . 90 

20. 19 

—  22.26 

25-33 

2.89 

6.93 

16.95 

5-98 

11.23 

15-19 

33-22 

4-57 

36.72 

21.38 

-   6.99 

4-38 

— 11.07 

7.66 

-   5.13 

0.  rubrinervis, igo5. 

Thickness  of  ovary 

Thickness  of  hypanthium 
Thickness  of  cone 

O.  gigas,  1905. 

Length  of  ovary 

Length  of  hypanthium. . . 

Length  of  cone ...    

Thickness  of  ovary 

Thickness  of  hypanthium 
Thickness  of  cone 

O.  lata,  1905. 

Length  of  ovary .     

Length  of  hypanthium. . . 

Length  of  cone 

Thickness  of  ovary 

Thickness  of  hypanthium 
Thickness  of  cone 


dM 

dV 

8  46 

II  .40 

14.84 

6.00 

10.26 

—   9.60 

9  52 

1.03 

20.67 

57-70 

1.54 

55-87 

37-03 

41 .  16 

42.38 

83-37 

29.63 

14.21 

10.53 

6.53 

15.06 

57-70 

22.21 

72  25 

18.13 

-^3-77 

13-87 

-    5.61 

13-87 

8.58 

In  this  table  all  the  deviations  which  are  less  than  the  sum  of  the  probable 
errors  are  printed  in  bold-face  figures,  and  may  be  left  out  of  the  discussion 
as  having  no  significance.  If  the  signs  of  the  remaining  numbers  are  taken 
into  account  it  is  found  that  16  of  the  30  characters  studied  showed  a  greater 
variability  in  the  mutant  than  in  the  parent,  while  only  4  were  significantly 
less  variable  in  the  mutant.  Thus  it  is  found  again,  as  in  1904,  that  there  is  a 
well-marked  tendency  for  the  mutant  to  show  a  greater  variability  than  its 
parent,  though  this  relation  is  far  from  being  absolute.  The  hint  then  recog- 
nized, that  the  degree  of  departure  from  the  mean  of  the  parent  might  be 
related  to  the  degree  of  increase  in  the  variability,  is  also  faintly  repeated,  but 
is  too  slight  to  be  now  considered  of  any  significance. 

The  single  study  in  the  correlation  of  characters  in  1904  showed  a  consid- 
erably less  correlation  in  the  mutant  than  in  the  parent  species,  and  we  have 
already  seen  in  the  study  of  leaf  characters  above  that,  though  that  study 
was  unsatisfactory,  such  evidence  as  there  is  seems  to  accord  with  the  view 
then  expressed  that  this  represents  a  general  fact.  In  order  to  gain  further 
evidence  on  this  point  the  coefficients  of  correlation  of  the  parts  of  the  buds 
were  calculated,  and  all  the  original  measurements  are  given  in  the  correlation 
tables  appended. 


36     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE)    OENOTHERAS. 


The  coefficients  of  correlation  are  as  follows : 


Length  of 
Length  of 
Length  of 
Length  of 
Length  of 
Thickness 
Length  of 
Length  of 
Length  of 
Thickness 
Length  of 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 


ovary 

hypanthium. 

ovary 

cone    


ovary 

of  ovary.  .  . 
hypanthium.  .  . 

cone 

hypanthium .     . 
of  hypanthium 

cone 

of  cone 

of  ovary 

of  hypanthium 

of  ovary 

of  cone 

of  hypanthium 
of  cone 


a 


!~    O 


-  o 

) 

) 


r  o 


r  o 


42i±o. 

563  ±o. 

477±o 

564±o. 

i57±o, 

873±o, 

558±o 

65i±o 

5i8±o 


044,0. 
0370. 
052  o. 
0370. 
052  o. 
0130. 
037,0. 
031  o. 
0390. 


387±o 
634±o 
600  ±0 
580  ±0 
044  ±0 
785±o 
573±o 
569  ±0 
532±o 


.0450 
.032  o 

.0340 
■0350 
•  0530 

.020  o 
.036  o 

.0360 

I 

.038;o 


472±o 
572±o 

272±0, 

535±o, 

i33±o. 

829±o, 

556±o. 

59i±o 

500  ±0 


041  o 

i 

0360 

049  o 

I 
0380 

052  o 

i 

017  o 

1 

i 

o37iO 
0350 
040  o 


298^0.049 
598±o.o34 
.596±o.o34 
375±o.046 
oi9±o.o5i 
835±o.oi6 
703±o.o27 
631  ±0.032 
,7i7±o.026 


Length  of 
Length  of 
Length  of 
Length  of 
Length  of 
Thickness 
Length  of 
Length  of 
Length  of 
Thickness 
Length  of 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 
Thickness 


ovary 

hypanthium . 


ovary . 
cone . . 


/ 


ovary 

of  ovary 

hypanthium.  . . 

cone 

hypanthium .  .  . 
of  hypanthium . 

cone 

of  cone 

of  ovary 

of  hypanthium . 

of  ovary 

of  cone 

of  hypanthium . 
of  cone 


0.22I  ±0. 
0.246±0. 

o.  336 ±0. 
o.507±o. 
o.i53±o. 
o . 840  ±  o . 
o.  5i4±o. 
o.5oi±o. 
o.540±o. 


0530 
052  o 
0500 

041  o 

I 

055I0 
o 


016 
042 
042 

035 


094±o 

■534±o 
.636±o 
527±o 
.438±o 
456±o 
83S±o 
475  ±0 
482+0 


0560 
.051  o 
.042  o 
.051  o 

•  0570 
•0560 

.  021  o 

.0750 
.054,0 


.624  ±0.043 
.638±o.o34 
.357. -to.  050 
.o25±o.o57 
.111  ±0.050 
837±o.oi7 
.64o±o.o34 
.584±o.o38 
.555±o.039 


A  comparison  of  the  coefficients  of  correlation  in  the  characters  of  the  sev- 
eral mutants  with  those  of  the  corresponding  characters  of  Oenothera  lama)xk- 
iana  shows  that  in  14  cases  out  of  27  there  is  a  decrease  of  correlation  in  the 
mutant ;  in  1 2  instances  there  is  an  increase,  and  in  i  the  correlation  remained 
the  same.  These  results  seem  to  negative  the  view  that  the  characters  of  the 
Oenothera  mutants  are  generally  less  closely  correlated  than  those  of  the  parent. 
However,  the  differences  between  the  coefficients  of  the  mutants  and  those 
of  Oenothera  lamarckiana  are  not  generally  greater  than  those  between  the 
several  lots  of  O.  lamarckiana,  or  between  the  two  lots  of  0.  riibrinervis,  and 
it  is  evident  that  the  question  is  still  to  be  considered  undecided,  but  with 
the  burden  of  evidence  in  favor  of  the  view  that  little  or  no  change  takes  place 
in  the  correlation  of  parts  when  mutation  changes  their  proportions. 


MUTATIONS,    VARIATIONS,    AND   REIvATlONSHIPS   OF   THE    OENOTHERAS.      37 


~" 

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Xo 

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Fig.  14. — Correlation  in  the  buds  of  Oenothera  lamarckiana , 
Xo.  04113.  Length  of  ovary  subject,  length  of  bud- 
cone  relative.  _CoefScient  of  correlation,  0.5720  ±  0.0358. 


1 

Xo 

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0    1 

2 

3 

4 

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6 

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2G  27  , 2.S  ]  29 :  TO 

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120 

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7 

7 

36 

24 

32 

23 

15 

10 

1 

0 

1 

160 

Fig.  15. — Correlation  in  the  buds  of  Oenothera  lamarckiana 
from  Hilversum.  Length  of  ovary  subject,  thickness  of 
ovary  relative.  Coefficient  of  correlation,  0.5997  i  0.0341. 


38     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


x„ 

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-4 

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Fig.  17. — Correlation  in  the  buds  oV Oeno- 
thera lamarckiana.  No.  04113.  Length 
of  ovary  subject,  thickness  of  ovary  rel- 
ative. Coefficient  of  correlation,  0.2724 
±0.0492. 


Fig.  16. — Correlation  in  the  buds  of  Oenothera  la- 
marckiana,'i^o.  04129.  Length  of  ovary  subject, 
thickness  of  ovary  relative.  CoeiBcient  of  cor- 
relation, 0.4766  ±0.0522. 


x. 

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_ 

Fig.  18. — Correlation  in  buds  of  Oenothera  lamarckiana  from  Hilversutn. 
Length  of  hypanthium  subject,  length  of  bud-cone  relative.  Coefficient  of 
correlation,  0.5797  ±  0.0354. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      39 


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3 

4 

5 

6 

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CO 

65 

70 

75 

80 

85 

90 

95  100 

105 

UO 

05 

70 

75 

80 

85 

90 

95 

lOO'lCS 

110!ll5 

Xi 

~  / 

32 

1 

2 

1 

4 

-6 

33 

2 

1 

3 

■~5 

31 

2 

2 

■ 

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35 

2 

4 

2 

8 

-3 

3C 

1 

G 

2 

9 

_2 

o7 

3 

10 

G 

3 

»>■> 

-1 

38 

8 

G 

4 

1 

19 

0 

39 

3 

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7 

1 

20 

1 

40 

6 

8 

2 

16 

0 

41 

3 

5 

3 

11 

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42 

1 

G 

9 

1 

17 

4 

43 

1 

5 

G 

5 

44 

4 

1 

5 

C 

45 

3 

3 

G 

7 

46 

2 

2 

4 

8 

47 

1 

2 

3 

9 

48 

1 

1 

1 

3 

1 

0    8 

36 

36 

34 

30 

9 

3 

0 

1 

158 

Fig.  24. — -Correlation  in  buds  of  O.  lamarcki- 
ana  from  Hilversum.  Lengtli  of  bud- 
cone  subject,  thickness  of  bud-cone 
relative.  Coefficient  of  correlation, 
0.7852 ±  0.0204. 


Fig.  25. — Correlation  in  the  buds  of  Oenothera 
lamarckiana.  No.  04129.  Length  of  bud-cone 
subject,  thickness  of  bud-cone  relative.  Coeffi- 
cient of  correlation,  0.8732  ±  0.0127. 


Xo 

-4 

f^ 

-2I-I 

0 

1 

2     3 

11 

C5 

70 

75 

SO 

S5 

90 

95  100 

70 

75 

80 

85 

90 

95 

'100105 

Xi 

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30 

1 

1 

-6 

31 

0 

-5 

32 

0 

-4 

33 

1 

1 

1 

3 

-3 

34 

1 

4 

5 

10 

-2 

35 

6 

12 

4 

1 

23 

-1 

36 

3 

6 

6 

2 

17 

0 

37 

1 

4 

13 

4 

22 

1 

38 

2 

13 

8 

1 

24 

2 

39 

1 

7 

10 

2 

1 

21 

— 

3 

40 

5 

9 

3 

17 

4 

41 

2 

4 

6 

5 

42 

5 

5 

6 

43 

1 

7 

1 

9 

7 

44 

1 

2 

3 

1 

2 

14 

32 

48 

38 

22 

4 

161 

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x. 

-4 

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0 

1 

2 

3 

4 

22 

23 

24 

25 

2G 

27 

28 

29 

30 

Xi 

—7 

2G 

1 

1 

-6 

27 

1 

1 

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2« 

1 

1 

-4 

29 

1 

1 

-3 

30 

1 

2 

3 

1 

7 

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31 

1 

1 

1 

3 

1 

7 

-1 

32 

2 

4 

11 

11 

5 

3 

36 

0 

33 

1 

3 

12 

G 

2 

24 

1 

34 

2 

6 

11 

11 

2 

32 

0 

35 

1 

1 

7 

13 

1 

23 

3 

30 

1 

2 

8 

3 

1 

15 

4 

37 

1 

1 

3 

5 

10 

5 

38 

1 

1 

6 

39 

0 

7 

40 

1 

1 

0 

6 

11 

27 

49 

47 

16 

1 

1 

160 

Fig.  26. — Correlation  in  the  bud  of  Oeno- 
thera lamarckiana.  No.  04113.  Length 
of  bud-cone  subject,  thickness  of  bud- 
cone  relative.  Coefficient  of  correla- 
tion, 0.8289  ±0.0166. 


Fig.  27. — Correlation  in  the  buds  of  Oenothera 
lamarckiana  from  Hilversum.    Thickness 
of  ovary  subject,  thickness  of  hypanthium . 
relative.    Coefficientof correlation, 0.5727 
±0.0358. 


42     MUTATIONS,    VARIATIONS,     AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


x. 

—  O 

-1 

0 

1 

2 

3 

4 

23 

24 

25 

26 

27 

28 

29 

Xi 

-5 

27 

1 

1 

-4 

28 

1 

3 

4 

-3 

29 

1 

1 

2 

_2 

30 

5 

5 

5 

2 

17 

-1 

31 

7 

4 

G 

4 

2 

23 

0 

32 

2 

G 

22 

8  1  3 

41 

1 

33 

O 

10 

21 

4 

37 

2 

34 

o 

id 

4 

3 

3 

12 

3 

35 

5 

1 

2 

2 

10 

4 

36 

1 

3 

1 

5 

5 

37 

1 

1 

2 

4 

6 

38 

1 

1 

2 

16 

24 

48 

47 

17 

3 

3 

158 

Fig.  28. — Correlation  in  the  bitds  of  O. 
lamarckiana,  0.4129.  Thickness  of 
ovary  subject,  thickness  of  hypan- 
thium  relative.  Coefficient  of  cor- 
relation, 0.5579  ±  0.0370. 


1 

Xo 

-3 

—o 

-1 

0 

1 

2 

3 

4 

5 

22 

23 

24 

25 

26 

27 

28 

29 

30 

Xi 

-3,28 

o 

3 

1 

1 

7 

—  n 

29 

1 

3 

3 

3 

6 

16 

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1 

4 

9 

13 

6 

2 

1 

36 

0 

31 

2 

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16 

5 

5 

34 

1 

32 

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5 

12 

10 

1 

30 

2 

33 

1 

9 

6 

4 

1 

1 

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3 

34 

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1 

1 

6 

10 

4 

35 

2 

1 

1 

4 

5 

36 

1 

1 

6 

37 

1 

1 

2 

12 

23 

49 

40 

22 

11 

1 

1 

161 

Fig.  29. — Correlation  in  the  buds  of  Oeno- 
thera lamarckiana,  Xo.  041 13.  Thickness 
of  ovary  subject,  thickness  of  hypan- 
thium  relative.  Coefficient  of  correlation, 
0.5565  +  0.0367. 


Xj-2 

-1 

0 

1 

2 

3|4 

5 

65 

70 

75 

80 

85 

90 

95 

100 

70 

75 

80 

85 

90 

95 

100 

105 

X, 

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2G 

1 

1 

-6 

27 

1 

1 

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1 

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10 

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5 

4 

3 

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4 

37 

3 

5 

1 

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10 

5 

38 

1 

1 

C 

39 

0 

7 

40 

1 

1 

5 

10 

39 

55 

38 

10 

2[1 

160 

Fig.  30. — Correlation  in  buds  of  Oenothera 
lamarckianairom  Hilversum.  Thick- 
ness of  ovary  subject,  thickness  of  bud- 
cone  relative.  Coefficient  of  correla- 
tion, 0.5691  ±  0.0360. 


Xo 

-4 

-3 

_o 

-1 

0 

1 

O 

3 

4 

5 

6 

60 

65 

70 

75 

80 

85 

90 

95 

100 

105 

110 

65 

70 

75 

80 

85 

90 

95 

lOOJlOS 

110115 

Xi 

-5  27 

1 

1 

-4|28 

4 

4 

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1 

1 

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1 

i 

S 

1 

17 

-1  31 

3 

11 

3 

5 

1 

09  1 

0 

32 

2 

7 

14 

14 

4 

41 

1 

33 

1 

2 

6 

8 

17 

o 

1 

37 

2  34 

3 

2 

2 

3 

2 

12 

3  35 

3 

4 

2 

1 

10 

4  ]36 

1 

1 

3 

5 

5  j37 

1 

2 

1 

4 

6 

38 

1 

1 

2 

1 

1 

0 

8 

36 

36 

34 

30 

9 

3 

0  1    158 

Fig.  31. — Correlation  in  the  buds  oi  Oenothera 
lamarckiana.  Xo.  04129.  Thickness  of  ovary 
subject,  thickness  of  bud-cone  relative. 
Coefficient  of  correlation,  0.6506  ±  0.0309. 


MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF    THE    OENOTHERAS.      43 


Xo 

L4 

■: 

-: 

-1 

0 

1 

0 

3 

05 

70 

7'> 

80 

85 

90 

95 

100 

TO 

75 

8U 

85 

00 

95 

100 

105 

X: 

-3 

2S 

0 

4 

1 

7 

—  0 

2y 

0 

3 

G 

5 

IG 

-1 

30 

1 

G 

12 

10 

I 

3G 

0 

31 

1 

14 

10 

i 

0 

34 

1 

32 

2 

2 

12 

7 

/ 

30 

0 

33 

S 

4 

5 

0 

00 

3 

3-1 

0 

G 

1 

10 

4 

35 

1 

0 

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4 

5 

30 

1 

1 

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37 

1 

1 

1 

•1 

14 

■il 

4S 

38 

00 

4 

IGl 

Xo 

-4 

-3 

_2 

-1 

0 

I 

2 

3     4     5 

G 

^^ 

60 

05 

70 

75 

SO 

85 

90 

95 

100 

105 

110 

05 

70 

75 

80 

85 

90 

95 

100 

105 

110 

115 

X. 

_2 

23 

1 

2 

y 

2 

t> 

16 

-1 

24 

2 

9 

5 

5 

3 

24 

0 

25 

3 

10 

15 

9 

10 

1 

4S 

1 

2G 

1 

G 

11 

15 

9 

4 

1 

47 

0 

27 

1 

0 

3 

G 

4 

1 

17 

3 

28 

1 

1 

1 

3 

4 

•29 

1 

1 

1 

3 

1 

0 

8 

3G 

3G 

34 

30 

9 

3 

0 

1 

158 

Fig.  32. —  Correlation  in  buds  of  O. 
lamarckiaiia.  No.  04113.  Thickness 
of  ovary  subject,  thickness  of  bud- 
cone  relative.  Coefficient  of  cor- 
relation.  0.5910  ±0.0346. 


Fig.  34. — Correlation  in  the  buds  of  Oenoiliera 
lamarckia.na,  No.  04129.  Thickness  of  hy- 
panthium  subject,  thickness  of  bud-cone  rela- 
tive. Coefficient  of  correlation,  0.5180 
±  0.0393. 


Xol-2 

-1 

0 

1 

0 

3 

4 

5 

65 

70 

75 

80 

85 

90 

95 

100 

70 

75 

80 

80 

90 

95 

100 

105 

Xi 

-4 

22 

1 

1 

0 

-3 

23 

1 

<> 

3 

G 

-2 

24 

3 

4 

0 

1 

1 

11 

-1 

25 

3 

9 

12 

3 

27 

0 

26 

1    17 

IG 

10 

3 

0 

49 

1 

27 

10 

15 

17 

5 

47 

2 

28 

6 

7 

2 

1 

IG 

3 

29 

1 

1 

i 

30 

1 

1 

5 

10 

39 

55  38 

10 

0 

1 

160 

1 

y^ 

-4 

-3 

_o 

-1 

0 

1 

2 

3 

65 

70 

75 

80 

85 

90 

95 

100 

70 

75 

SO 

85 

90 

95  100 

105 

X: 

_3 

00 

1 

1 

•-> 

_o 

23 

5 

G 

1 

12 

-1 

24 

1 

4 

7 

i 

4 

23 

0 

25 

1 

2 

9 

17 

13 

/ 

49 

1 

2G 

1 

7 

10 

14 

7 

1 

40 

2 

27 

1 

3 

9 

5 

3 

1 

22 

3 

28 

2 

2 

5 

0 

11 

4 

29 

1 

1 

•1 

30 

1 

1 

I 

2 

14 

32 

48  38 

22 

4 

IGl 

. 

Fig.  33. — Correlation  in  buds  of  O.  lam- 
arckiana  from  Hilversum.  Thick- 
ness of  hypanthium  subject,  thick- 
ness of  bud -cone  relative.  Coefficient 
of  correlation,  0.5316  ±  0.0382. 


Fig.  35. — Correlation  in  the  buds  of  O. 
lamarckiaiia.  No.  04113.  Thickness 
of  hypanthium  subject,  thickness 
of  bud-cone  relative.  Coefficient 
of  correlation,  0.5005   ±0.0398. 


Xo 

-9 

-8 

-7 

-6 

-5 

-4 

-3 

-2 

-1 

0 

1 

2 

3 

4 

5 

6 

7 

8 

9 

21  i  22 

23 

24 

25 

26 

27 '28  29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

X. 

-4 

85 

90 

1 

2 

1 

1 

5 

-3 

90 

95 

2 

1 

3 

2 

2 

1 

11 

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95 

100 

1 

1 

2 

1 

1 

1 

1 

1 

2 

1 

2 

2 

16 

-1 

1001C5 

1 

2 

4 

1 

4 

ty 

4 

8 

2 

1 

2 

1 

1 

33 

0 

105110 

1 

5 

2 

3 

1 

3 

3 

3 

1 

2 

1 

1 

1 

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1  110115 

1 

2 

1 

1 

2 

1 

5 

3 

1 

2 

2 

4 

3 

1 

1 

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1 

1 

1 

2 

1 

1 

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125,130 

1 

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2 

2 

3 

4 

8 

8 

13 

10 

13 

15 

20 

18 

7 

13 

7 

7 

6 

2 

0 

IGO 

Fig.  36. — Correlation  in  the  buds  of  Oenothera  rubrinervis.  No.  04113. 
Length  of  ovary  subject,  length  of  hypanthium  relative.  Co- 
efficient of  correlation,  0  .  2985  ±  0.0486. 


44     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS. 


' 

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12 

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14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35  "si? 

37 

X, 

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95 

1 

1 

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95 

100 

1 

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100 

105 

1 

1 

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105 

110 

1 

2 

2 

1 

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1 

1 

10 

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110 

115 

4 

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4 

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1 

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120 

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125 

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110 

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15 

15 

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Fig.   37. — Correlation  in  the  buds  of  Oenothera  ruhrincrvis,  Xo.'04137.      Length  of  ovary  subject,  length 
of  hypanthium  relative.     Coefficient  of  correlation,  0.2211  ±  0.0531. 


i 

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11 

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27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

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X: 

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85 

90 

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95 

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1 

2 

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100 

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105 

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105 

110 

1 

1 

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5 

4 

3 

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1 

110 

115 

1 

1 

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6 

0 

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120 

1 

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125 

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130 

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3 

4 

i 

9 

12 

16 

21 

24 

19 

17 

8 

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5 

2 

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Fig.   38. — Correlation  in  the  buds  of   Oenothera  rubrinervis,  Xo.  04113.      Length  of  ovary  subject,  length 
of  bud-cone  relative.     Coefficient  of  correlation,  0.5976  ±  0.0343. 


~ 

~ 

"~ 

"-^ 

X. 

-11 

40 

-9 

-8 

-7 

-6 

-5 

-4 

-3 

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1 

2 

3 

4 

5 

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25 

26 

27 

28 

29 

30 

31 

32 

33 

31 

35 

3G 

37 

3s 

39 

40 

41 

42 

Xi 

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90 

95 

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95 

100 

1 

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100 

105 

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no 

1 

2 

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1 

1 

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110 

115 

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120 

125 

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125130 

1 

3 

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133 

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12 

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0 

4 

3 

22 

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27 

23 

21 

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4 

3 

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146 

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Fig.  39. — Correlation  in  the  buds  of  Oenothera  rubrinenns.  No.  04137.     Length  of  ovary  subject,  length 
of  bud-cone  relative.     Coefficient  of  correlation,  0.2464  ±  0.0524. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS.      45 


- 

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Fig.  40. — Correlation  in  the  buds  of  Oenothera  ru- 
brinenis,  No.  04113.  Length  of  ovary  subject, 
thickness  of  ovary  relative.  Coefficient  of  correla- 
tion, 0 .  5957  ±  0 .  0344. 


Fig.  41. — Correlation  in  the  buds  of  O.  ruhrinervis , 
Xo.  04137.  Length  of  ovary  subject,  thickness 
of  ovary  relative.  Coefficient  of  correlation, 
0.3357  ±0.0495. 


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Fig.  42. — Correlation  in  the  buds  of  Oenothera  rubrinervis.tio.  0-i\\3.  Length 
of  hypanthium  subject,  length  of  bud-cone  relative.  Coefficient  of  correla- 
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46     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 


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48     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS. 


X, 

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Fig.  48. — Correlation  in  the  buds  of  Oenothcrj, 
rubrinenis.  No.  04113.  Thicknessof  ovary 
subject,  thickness  of  hypanthium  relative. 
CoefiBcient  of  correlation,  0 .  7026  ±  0 .  0270.- 


1 

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146 

Fig.  49. — Correlation  in  the  buds  of 
Oenothera  rubrinervis,  No.  04137. 
Thickness  of  ovary  subject,  thick- 
ness of  hypanthium  relative.  Coef- 
ficient of  correlation,  0 .  504 1  ±  0. 04 1 6. 


Xo 

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1 

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— 

1 

Fig.  50. — Correlation  in  the  buds  of  Oeno- 
thera rubrinervis, 'Ho.  04113.  Thickness 
of  ovary  subject,  thickness  of  cone  rel- 
ative. Coefficient  of  correlation,  0.6310 
±  0.0321. 


IXnl-4!-S 

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0 

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43 

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'1 

146 

Fio.  51. — Correlation  in  the  buds  of  Oeno- 
thera rubrinervis,  Xo.  04137.  Thick- 
ness of  ovary  subject,  thickness  of 
bud-cone  relative.  Coefficient  of  cor- 
relation, 0.5006  ±0.0418. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE)    OENOTHERAS.     49 


X2-4 

-3 

_o 

-1 

0 

1 

0 

3 

-> 

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75 

80 

85 

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95 

100 

105 

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90 

95 

100 

105 

110 

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5 

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Fxo.  52. — Correlation  in  buds  of  O.  rubri- 
nei~iis,^o.  04113.  Thickness  of  hypan- 
thium  subject,  thickness  of  bud-cone 
relative.  Coefficient  of  correlation, 
0.7170  ±0.0259. 


Fig.  53. — Correlation  in  the  buds  of 
O.  rtibrin-ertis.  No.  04137.  Thickness 
of  hypanthium  subject,  thickness  of 
bud-cone  relative.  Coefficient  of  cor- 
relation, 0  .5397  ±  0.0354. 


Xo 

-9 

-8 

-7 

-6 

-5 

-4 

-3 

-2 

-1 

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Fig.   54. — Correlation    in    the  buds  of  Oenothera  gigas.      Length   of   ovary  subject,   length 
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Fig.  55. — Correlation  in  the  buds   of  O.  gigas.      Length   of  ovary  subject,  length 
of  bud-cone  relative.      Coefficient  of  correlation,  0.5338  ±  0.0506. 


50     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   O^   THE    OENOTHERAS. 


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Fig.  56. — Correlation  in  the  buds  of  Oenothera  gigas.  Length  of 
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relation,   0.8385  ±0.0210. 


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Fig.  61. — Correlation  in  the  buds  of  Oenothera  gigas. 
Thicknessof  ovary  subject,  thickness  of  bud-cone 
relative.  Coefficient  of  correlation,  0.4750  ± 
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Fig.  62. — Correlation  in  the  buds  of  Oenothera  gigas. 
Thickness  of  hypanthium  subject,  thickness  of 
bud-cone  relative-  Coefficient  of  correlation, 
0.4822  ±0.0543. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      5;^ 


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Fig.   63. — Correlation  in  the  buds  of  Oenothera   lata.    Length   of  ovary  subject,  length  of 
hypanthium  relative.     Coefficient  of  correlation,  0.0944  ±  0.0565. 


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Fig.   64. — Correlation  in  the  buds  of  Oenothera  lata.     Length  of  ovary  subject,  length 
of   bud-cone   relative.     CoefiScient  of  correlation,  0.6382  rb  0.0338. 


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Fio.  65. — Correlation  in  the  buds  of  Oenoiliera 
lata.  Length  of  ovary  subject,  thickness 
of  ovary  relative.  Coefi&cient  of  correla- 
tion, 0.3568  ±0.0497. 


54     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


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MUTATIONS,    VARIATIONS,    AND   REI.ATIONSHIPS   OF    THE    OENOTHERAS.      55 


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Fig.  68. — Correlation  in  the  buds  of  Oenottiera 
lata.  Length  of  bud-cone  subject,  thickness 
of  bud-cone  relative.  CoefiBcient  of  correlation, 
0.8366  ±0.0171. 


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Fig.  69. — Correlation  in  the  buds  of  Oen- 
othera lata.  Thickness  of  ovary  sub- 
ject, thickness  of  hypanthium  rela- 
tive. Coefficient  of  correlation,  0 .  6400 
±0.0336 


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Fig.  70. — Correlation  in  the  buds  of  Oenothera 
lata.  Thickness  of  ovary  subject,  thickness 
of  bud-cone  relative.  Coefficient  of  corre- 
lation, 0.5839  ±0.0376. 


Fig.  71. — Correlation  in  the  buds  of  Oenothera 
lata.  Thickness  of  hypanthium  subject, 
thickness  of  bud-cone  relative.  Coefficient 
of  Icorrelation,  0.5549  ±  0.0394. 


56     MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF   THE    OENOTHERAS. 

PEDIGREE-CULTURES  OF  OENOTHERA  GRANDIFLORA,  O.  BIENNIS,  AND 

O.  CRUCIATA. 

In  order  to  test  the  mutative  capacity  of  other  evening-primroses,  seeds 
were  obtained  from  correspondents  in  various  parts  of  the  world  and  as  many 
as  40  sowings  were  made  in  one  year  from  0.  biennis.  In  only  one  strain, 
however,  did  anything  come  to  light  which  would  be  of  interest  in  connection 
wnth  the  chief  purpose  of  these  investigations. 

Parental  individuals  were  selected  and  verified  by  Dr.  N.  L.  Britton  in  1903, 
and  from  the  seeds  furnished  by  them  the  plants  were  grown  which  furnished 
material  for  the  descriptive  diagnosis  published  in  a  previous  paper  (Mac- 
Dougal,  Vail,  ShuU  &  Small,  1905).  This  is  not  the  species  growing  wild  in 
Europe  and  cited  by  De  Vries  in  his  "Mutationstheorie. " 

Inflorescences  of  4  separate  individuals  were  inclosed  in  transparent  paper 
bags  in  such  manner  as  to  secure  pure  fertilization  and  the  seeds  preserved  in 
separate  packages.  Sowings  were  made  in  pans  of  sterilized  soil,  in  accord- 
ance with  the  usual  custom,  early  in  January,  1905. 

A  number  of  seedlings  were  seen  in  which  the  first  foliage-leaf  and  all  suc- 
ceeding leaves  were  very  much  narrower  than  the  parental  type.  Sixteen 
individuals  of  this  kind  were  found  and  preserved,  appearing  in  the  progeny 
of  all  of  the  4  parents  represented.  An  exact  count  of  the  number  was  kept 
only  in  the  case  of  one  parent.  No.  8.17,  in  which  4  of  these  aberrants  were 
included  in  a  progeny  of  669  individuals  and  formed  very  nearly  0.6  per  cent 
of  the  entire  number. 

The  aberrants  retained  their  property  of  forming  narrow  leaves,  which  w'ere 
slightly  paler  in  color  than  the  parental  type,  and  appeared  to  taper  gradually 
at  the  base  into  long  petioles.  One  aberrant  of  8.17  gave  off  a  lateral  branch 
at  the  base  of  the  stem  which  developed  broader  leaves  resembling  those  of 
the  parent.  With  the  completion  of  development,  the  form  of  the  capsules 
and  the  entire  habit  of  this  branch  were  in  exact  duplication  of  O.  biennis. 

On  July  2  one  of  the  aberrants  opened  a  flower  which  also  showed  some 
departures  from  the  normal,  being  characterized  by  the  extremely  long  sta- 
mens and  by  the  elongated  capsule,  which  was  of  equal  diameter  throughout 
and  did  not  taper  to  a  point  or  toward  the  tip  as  in  its  parental  form.  The 
first  4  flowers  which  were  brought  to  maturity  on  a  lateral  branch  were  inclosed 
in  a  bag  and  self -pollinated.  The  seeds  formed  were  allowed  to  mature  and 
these  were  sown  as  soon  as  practicable  in  order  to  determine  the  constancy 
of  the  new  form. 

The  adult  plant  has  a  main  stem  varying  from  80  to  150  cm.  in  height,  and 
branches  freely  from  the  base  upward.  The  lower  branches  are  long,  slender 
and  assurgent,  while  the  upper  ones  are  much  shorter.     Numerous  secondary 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      57 

branches  arise  during  the  latter  part  of  the  season.     The  stems  are  nearly 
terete  below,  but  are  slightly  channeled  above. 

The  stem-leaves  are  10  to  16  cm.  long,  and  not  more  than  i  cm.  in  width 
at  the  middle,  obscurely  pubescent  on  both  surfaces,  thickish,  yellowish  green, 
obscurely  and  irregularly  denticulate,  with  a  heavy,  broad  midvein.  The 
leaves  are  linear,  and  taper  to  the  long  acuminate  tip  and  to  the  margined 
petiole. 

The  bracts  are  narrowly  linear  and  3  cm.  long.  The  conic  portion  of  the 
bud  is  about  12  mm.  long  and  half  that  in  diameter,  pale  yellow,  and  barely 
tapering  in  the  apical  portion,  being  finely  pubescent.  The  free  erect  tips  are 
irregularly  acuminate  and  2  to  4  mm.  long. 

The  hypanthium  is  25  to  35  mm.  long  and  exceeds  the  cal^TC-lobes  by  half. 

The  filaments  are  7  to  9  mm.  long,  stout,  and  the  anthers  4  mm.,  slender. 
The  pistil  is  slightly  shorter  or  longer  than  the  stamens,  measuring  4  to  6 
mm.,  and  this  species  is  therefore  self-pollinated  like  the  parental  form.  The 
stigmatic  lobes  are  4  to  5  mm.  long  and  spreading. 

The  capsules  are  25  to  35  mm.  long,  4  to  5  mm.  in  diameter,  generally 
curv^ed,  sparingly  pubescent,  bright  green  and  shining,  scarcely  tapering  to  the 
apex,  and,  in  general,  much  thinner  than  the  parental  type.  The  ovary  is 
about  half  or  less  than  half  the  length  of  the  mature  capsule. 

The  sepals  are  3  mm.  long,  the  petals  firm,  15  to  16  mm.  long  and  15  mm. 
wide,  being  crenately  emarginate  (1905  c,  p.  17). 

A  small  package  of  the  lot  of  seeds  from  which  the  mutant  was  obtained 
originally  were  sent  to  INIiss  Elizabeth  Billings,  who  sowed  them  in  her  green- 
house at  Woodstock,  Vermont,  in  March,  1905,  and  later  transferred  them  to 
her  experimental  garden.  The  499  plants  thus  obtained  were  kept  under 
observation  by  Miss  Billings  and  Professor  F.  S.  Lee,  of  Columbia  University, 
with  the  result  that  aberrant  individuals  were  found  which  were  identical 
with  those  obtained  during  the  previous  September  in  the  New  York  Botanical 
Garden. 

The  development  of  the  newly  arisen  form  has  been  followed  from  Septem- 
ber, 1904,  until  the  present  time.  The  first  individual  discovered  came  into 
bloom  on  July  2,  1905,  having  been  cultivated  under  glass  during  the  preced- 
ing Mdnter.  Seeds  were  matured  5  or  6  wrecks  later,  and  sowings  have  been 
made  from  them,  with  the  result  that  the  aberrant  type  was  found  to  produce 
a  progeny  which  contained  not  more  than  12  per  cent  of  the  atypic  form  of 
individuals,  the  remainder  being  of  the  parental  type  and  unchanged.  Hybrid- 
izations between  the  aberrant  form  and  the  parental  form  gave  the  same 
result,  while  the  aberrant  type  when  crossed  with  lamarckiana  and  with  ruh- 
rinervis  gave  a  progeny  resembling  in  general  constituency  those  produced 
by  typical  biennis. 


58     MUTATIONS,   VARIATIONS,    AND  RELATIONSHIPS   OF  THE   OENOTHERAS. 

A  similar  aberrant  of  0.  cniciata  has  been  found  in  many  of  the  cultures  of 
that  species.  Much  attention  has  been  devoted  to  these  forms,  both  as  to 
habits  of  growth  and  anatomy.  The  only  parallel  occurrences  are  those 
which  are  to  be  found  in  other  genera  in  which  atypical  individuals  affected  by 
the  enzymatic  diseases  may  give  a  progeny  in  which  the  normal  form  of  indi- 
viduals predominates.  The  vigorous  growth  of  the  atypic  individuals  and 
the  comparative  regularity  with  which  it  appears  are  well  worthy  of  remark. 
If  these  forms  were  found  growing  in  the  open,  they  might  easily  be  taken  as 
belonging  to  a  species  apart  from  0.  biennis  unless  their  progeny  was  tested 
in  pure  cultures. 

The  pedigreed  cultures  of  0.  grandiflora  made  in  1906  from  purely  fertilized 
seeds  of  the  previous  year  included  about  1 500  plants.  More  than  one  season 
of  such  cultures  is  necessary  to  secure  definite  results  that  may  be  depended 
upon.  The  observations  have  been  carried  so  far  at  the  New  York  Botan- 
ical Garden  and  the  Desert  Laboratory,  however,  as  to  warrant  the  assertion 
that  this  species  presents  a  complex  progeny  analogous  to  that  of  lamarckiana, 
in  which  two  well-defined  mutants  are  readily  recognizable  by  reason  of  their 
striking  diflferences  from  the  parental  form. 

ORIGIN  OF  FIXED  FORMS  BY  HYBRIDIZATION. 

That  the  cross-fertilization  of  two  forms  may  result  in  the  production  of  a 
unitypic  or  polytypic  progeny  in  which  the  parental  qualities  appear  in  a 
mosaic  is  well  known.  Components  of  a  native  flora  have  been  suspected  to 
be  of  hybrid  origin  in  a  few  instances,  and  by  succeeding  experimental  tests 
have  been  synthetized  from  the  parents.  The  practice  prevailing  among 
taxonomists  of  ascribing  a  hybrid  origin  to  a  newly  discovered  form,  which, 
in  outward  anatomical  characters,  is  between  two  know^n  species,  is  extremely 
pernicious  and  is  not  justified  by  facts  obtained  in  cultural  work.  The  best 
grounds  for  such  a  conclusion  are  to  be  found  when  two  species  alone  occur 
in  a  region,  and  the  appearance  of  a  third  is  attributed  to  hybridization ;  but 
even  here  the  supposition  that  a  mutation  may  have  ensued  is  allowable. 

The  hybrid  0.  lamarckiana  X  0.  cruciata  (form  with  long  hypanthium  and 
slender  bud),  which  was  described  in  the  previous  publication  by  the  authors, 
proves  to  be  a  fixed  form,  and  as  it  sets  seeds  freely  and  is  self-fertilizing,  it  is 
in  every  respect  an  independent  species.  Several  hundreds  of  seedlings  were 
grown  from  purely  fertilized  capsules  ripened  in  1904,  with  the  result  that  all 
w^ere  seen  to  conform  to  the  parental  hybrid  type  in  every  particular.  The 
two  parental  forms  do  not  meet  in  their  native  habitats,  and  hence  this  form 
could  not  have  arisen  in  a  state  of  nature. 

The  reciprocal  of  this  cross,  that  is,  0.  cruciata  X  0.  lamarckiana,  consists  of 
three  forms,  one  of  which  is  indistinguishable  from  the  0.  biennis  cruciata 
which  has  been  received  from  various  European  gardens. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF  THE   OENOTHERAS.     59 

BUD-SPORTS— VEGETATIVE  SALTATIONS. 

One  of  the  most  interesting  and  least  understood  phases  of  heredity  is  that 
in  which  the  tract  of  embryonic  tissue  constituting  a  bud  may  develop  and 
put  into  external  realization  a  set  of  characters  wholly  different  from  those  of 
the  remainder  of  the  individual.  A  large  number  of  such  instances  have  been 
seen  by  gardeners,  horticulturists,  and  farmers,  although  exact  observations 
on  the  hereditary  qualities  of  such  mutant  branches  are  almost  wholly  want- 
ing. In  some  cases  the  departure  from  the  type  does  not  affect  the  entire 
branch,  and  it  may  not  be  made  manifest  until  an  advanced  stage  of  its  devel- 
opment, causing  some  of  the  leaves  or  perhaps  some  of  the  flowers  to  show 
atypic  characters.  When  such  partial  vegetative  saltation  ensues  it  gen- 
erally results  in  converting  a  lateral  section  of  the  branch  or  inflorescence  into 
the  new  form,  and  has  been  termed  "sectorial  variation"  byDeVries.  To 
designate  it  beyond  danger  of  misapprehension,  however,  it  should  be  known 
as  sectorial  bud-mutation. 

The  cultures  carried  on  in  the  experimental  garden  during  1905  were  char- 
acterized by  two  remarkable  bud-sports. 

The  hybrid  0.  lamarckiana  X  {0.  lamarckiana  X  0.  cruciata)  comprised 
nearly  a  score  of  recognizable  forms  in  the  seedling  stage.  About  two  of 
each  type  were  transplanted  to  the  experimental  garden  in  May,  1905.  Two 
of  the  types  included  were  apparently  different  from  each  other  only  in  the 
character  that  one  of  them  bore  flowers  with  broad  petals  resembling  those  of 
lamarckiana,  while  the  other  was  furnished  with  cruciate  flowers,  but  with  the 
petals  broader  than  the  typical  cruciata  and  quite  as  long  as  those  of  lamarck- 
iana. It  was  evident  that  the  two  forms  constituted  an  illustration  of 
Mendelian  combinations,  alike  in  all  particulars  except  as  to  the  characters 
imited  in  the  form  and  size  of  the  petals. 

One  of  the  individuals  with  the  broadly  cruciate  flowers  bore  a  branch  near 
the  base  of  the  stem,  on  which  were  formed  only  flowers  of  the  broadly-petaled 
lamarckiana  type.  Some  of  these  were  purely  fertilized  and  seeds  preserved 
for  testing.  These  were  duly  sown  in  December,  1905,  under  glass,  and 
representative  individuals  brought  to  bloom  in  the  open  air  in  July,  1906. 

When  they  came  into  bloom  in  July,  1906,  it  was  found  that  the  branch- 
sport  came  entirely  true  to  the  type  which  it  represented.  With  this  it  is 
interesting  to  note  that  the  purely  fertihzed  seeds  of  the  main  stem  bearing 
cruciate  flowers  gave  a  progeny  which  contained  some  individuals  bearing 
flowers  with  broad  petals,  while  the  other  strain,  which  bore  only  broad 
petals,  came  entirely  true  to  its  type  without  deviation.  No  exact  numerical 
count  was  made,  but  it  seemed  evident  that  the  cruciate  petal  is  dominant 
over  the  cordate,  and  that  the  plants  described  above  exemplify  the  develop- 
ment of  a  sport  on  a  plant  of  the  first  generation  in  which  the  recessive  char- 
acter appeared  and  when  extracted  came  true  to  the  recessive  character. 


6o     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

De  Vries  (1900,  p.  86)  described  a  case  of  vegetative  mutation  in  a  hybrid 
between  Veronica  longifolia,  which  has  a  blue  flower,  and  V .  alba  with  a 
white  flower.  The  hybrid  has  a  blue  flower,  but  several  cases  were  noted  in 
which  the  entire  buds  or  portions  of  inflorescences  produced  white  flowers. 
These  white  sports,  when  self-fertilized,  produced  white  flowers  only,  in  the 
preliminary  tests  cited. 

Similar  vegetative  splitting  is  known  in  Datura  hybrids,  being  seen  by 
Naudin  in  3  individuals  only,  while  sports  of  this  character  of  Brassica,  Raph- 
aniis,  Abies,  Anagallis,  Hclianthemum,  Zea,  and  Cytisiis  are  well  known. 
Nearly  all  other  sports  of  hybrids  are  clearly  of  an  atavistic  nature,  being 
more  or  less  direct  reversions,  sometimes  rehearsing  the  juvenile  characters 
of  one  of  the  parents  of  the  hybrids. 

A  second  instance  was  offered  by  a  form  known  as  0.  ammophila.  This 
material  was  grown  from  seeds  obtained  from  Professor  W.  O.  Focke,  which 
had  been  collected  on  the  coast  near  Bremen,  Germany. 

A  large  number  of  seeds  were  sown  in  sterilized  soil  in  the  propagating 
house  on  February  10,  1905.  A  month  later  the  seedlings  obtained  were 
examined  and  a  dozen  representing  the  widest  visible  range  of  variation  were 
transferred  to  pots,  while  the  remainder  of  the  culture  was  discarded.  With 
successive  repottings  the  number  of  individuals  was  reduced  to  7,  which  were 
transplanted  to  the  experimental  garden  in  the  latter  part  of  May,  at  which 
time  no  differences  of  moment  were  noticed.  During  the  summer  3  of  the 
specimens  sent  up  shoots,  while  the  other  4  formed  dense  rosettes  with  a  few 
short  lateral  branches. 

About  the  middle  of  August  the  more  advanced  individuals  came  into 
bloom.  No  careful  examination  of  them  had  been  made  up  to  this  time, 
but  it  was  now  seen  that  one  of  them  had  formed  a  lateral  branch  at  the 
base  of  the  main  stem,  which,  by  the  spread  of  its  branches,  the  shape  and 
expansion  of  its  leaves,  and  its  general  vigorous  growth  had  become  the  larger 
member  of  the  shoot.  This  branch,  in  the  form  and  behavior  of  all  of  its 
organs,  including  flowers  and  fruits,  was  an  exact  reproduction  of  the  O. 
biennis  which  has  formed  the  basis  of  cultures  under  that  name  in  the  New 
York  Botanical  Garden.  Branches  of  the  sport  and  of  typical  biennis  were 
submitted  to  several  botanists,  with  the  result  that  they  were  found  to  be 
indistinguishable  by  anatomical  characters. 

The  main  stem,  which  had  branched  in  the  usual  manner,  had  been  crowded 
from  its  natural  upright  position  and  had  assumed  a  half- recumbent  position, 
having  the  appearance  of  a  lateral  branch. 

The  suggestion  lies  near  at  hand,  that  O.  ammophila  is  a  hybrid  derivative 
of  O.  biennis,  and  that  the  vegetative  mutation  is  simply  one  of  reversion,  after 
the  manner  of  examples  cited  above.  No  positive  evidence  upon  the  origin  of 
ammophila  is  at  hand,  however. 


o. 


PLATE  9. 


Ot-nothera  aminophila  with  bud-sport.  The  narrow -leaved  decumbent  stock  represents  the  typical  parent, 
while  the  more  vigorous  upright  branch  with  broader  leaves  is  the  mutant  branch  which  represents 
Oenothera  biennis. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      6 1 

Purely  fertilized  seeds  of  the  mutant  branch  and  of  the  main  stem  were 
obtained,  and  these  were  sown  late  in  September,  1905.  Those  of  the  main 
stem  came  true  to  the  type  of  the  main  stem,  that  is,  to  0.  ammophila.  All 
of  the  progeny  obtained  from  the  purely  guarded  seeds  of  the  sport  conformed 
strictly  to  the  biennis  type  (plate  9). 

THE  INDUCTION  OF  MUTATIONS. 

One  of  the  most  important  problems  confronting  the  investigation  in  this 
subject  is  that  of  the  localization  of  the  mutations  in  the  Hfe-history  of  the 
plant.  If  we  could  fix  upon  the  exact  stage  in  which  the  direct  changes 
occurred,  we  then  would  be  in  a  position  to  examine  the  protoplasts  con- 
cerned with  a  view  to  ascertaining  what  changes  ensue  in  the  chromosomes  in 
connection  with  saltations  in  inheritance.  Furthermore,  opportunity  would 
also  be  offered  for  attempts  to  determine  the  factors  which  operate  as  stim- 
uli, or  loosing  agents  in  setting  the  mutations  free. 

A  theoretical  consideration  of  the  subject  seemed  to  indicate  that  the 
changes  constituting  the  mutations  which  give  rise  to  atypic  seedlings  took 
place  in  a  stage  previous  to  the  reduction  divisions  in  the  embryo-sac  or  in 
the  pollen  mother-cell.  It  was  planned,  therefore,  to  subject  these  structures 
to  the  action  of  chemical  agents  at  a  time  before  fertilization  had  occurred. 
The  attempt  was  made  to  secure  two  forms  of  stimulation  by  using  some  solu- 
tions of  high  osmotic  value,  and  other  mineral  compounds  which  are  stimu- 
lative in  low  concentrations.  A  desideratum  in  such  experimentation  was  to 
use  plants  in  which  a  large  number  of  ovules  were  to  be  found  in  one  ovarial 
cavity.  The  solutions  were  injected  into  the  ovaries  by  means  of  a  physician's 
hypodermic  syringe.  Operations  of  this  character  were  carried  out  with 
Begonia  rotundifolia  and  with  a  species  of  Cleome,  but  with  negative  results, 
and  injections  were  also  made  into  the  central  placenta  of  a  number  of  flowers 
of  Abutilon  abutilon,  but  in  all  cases  at  a  very  late  stage,  and  so  far  no  results 
have  been  obtained  with  this  plant.  It  is  being  used  in  a  new  series  of  experi- 
ments, however,  which,  with  improved  technique,  may  be  expected  to  furnish 
some  interesting  material. 

Attention  was  next  turned  to  several  species  of  Oenothera  which  were 
being  cultivated  in  pure  pedigreed  strains  in  the  experimental  garden. 
Solutions  of  copper  sulphate  of  i  to  400,000  were  injected  into  ovaries  of 
lamarckiana  immediately  previous  to  pollination  and  the  pistils  were  purely 
pollinated.  The  capsules  formed  from  ovaries  treated  in  this  way  were  vari- 
ously distorted  as  a  result  of  the  wounding,  but  a  large  crop  of  perfect  seeds 
was  matured,  which  were  sowed  in  September,  1905.  A  careful  census  of  the 
seedlings  late  in  November  made  it  appear  that  in  the  small  number  of  mutants 
present  the  usual  mutants  of  this  species  occurred  in  normal  proportions. 


62     MUTATIONS,    VARIATIONS,    AN'D   RELATIONSHIPS    OF   THE    OENOTHERAS. 

A  number  of  radium  pencils  were  taken  from  a  series  used  by  Dr.  Gager  in 
his  investigations,  and  were  fastened  in  an  inflorescence  in  such  manner  that  a 
deleterious  influence  must  have  been  exerted  on  some  ovules,  while  in  other 
cases  the  effect  must  have  been  stimulative  if  the  reactions  of  seedlings  of 
other  plants  may  be  taken  to  offer  a  fair  analogy.  The  crop  of  seeds  matured 
in  the  treated  ovaries  was  sown  in  September,  1905,  and  late  in  November  a 
census  showed  that  only  20  normal  specimens  of  laniarckiana  were  present 
and  that  2  mutants  were  also  included.  The  rate  of  mortality  was  thus  seen 
to  be  twice  as  great  in  the  type  as  in  the  mutants  in  this  single  test. 

Similar  tests  were  made  with  0.  biennis.  Solutions  of  magnesium  sulphate 
in  distilled  water  were  used  without  any  noticeable  departure  in  the  composi- 
tion or  behavior  of  the  progeny  arising  from  the  seeds.  Poisonous  solutions 
containing  i  part  of  zinc  sulphate  in  500  of  distilled  water  were  also  used. 
The  seeds  produced  in  the  ovaries  which  had  received  this  treatment  contained 
atypic  forms  of  the  kind  seen  in  ordinary  cultures  in  apparently  the  cus- 
tomary proportion  of  about  one  in  200.  In  addition  however,  a  single  rosette 
was  found  which  differed  widely  from  any  known  type,  and  of  this  form,  which 
was  recognizably  different  from  the  parental  type  in  many  qualities,  some  of 
the  differences  were  plainly  apparent  even  in  the  earliest  leaves  of  the  seedlings. 
These  differences  have  become  accentuated  in  the  adult  plant. 

The  parental  form  has  been  under  observation  for  five  years  in  cultures 
and  in  a  wild  condition.  An  aberrant  form,  which  appears  to  be  eversporting, 
has  been  previously  figured,  and  while  this  form  appeared  in  the  injected 
or  treated  seeds  in  a  normal  proportion,  yet  the  newest  aberrant  has  not  been 
seen  elsewhere.  The  probability  must  be  taken  into  account  that  it  may  be 
a  mutant  of  rare  occurrence,  the  cycle  of  which  came  within  the  experiments ; 
but  in  either  case  it  is  plainly  a  mutant,  and  it  only  remains  to  be  seen  whether 
or  not  it  was  induced  by  the  action  of  the  zinc  solution.  The  presumption 
seems  to  favor  such  a  conclusion.  Seeds  of  this  form  were  harvested  in 
August,  1906,  and  sown  immediately,  with  the  result  that  the  characters  of 
the  new  form  were  found  to  be  fully  transmissible,  the  first  generation  of  the 
progeny  being  duplicates  of  the  parent  within  the  limits  of  fluctuating  varia- 
bility. The  entire  plant  is  characterized  by  a  much  deeper  green  color  than 
the  parental  form  and  the  leaves  are  slightly  curled  and  twisted,  owing  to 
inequalities  of  growth,  and  it  reaches  maturity  quite  early  in  the  season. 

In  addition  to  the  crop  of  guarded  seeds  harvested  from  some  of  the 
branches,  the  available  remainder  were  collected  and  sown  in  the  greenhouse  of 
the  Dessrt  Laboratory,  with  the  result  that  this  second  generation  of  the  deriv- 
ative was  found  to  conform  exactly  in  every  individual  to  the  derivative  type. 
Either  the  mutant  does  not  intercross  with  the  parent,  although  the  branches 
were  in  contact,  or  if  a  cross  has  occurred  the  derivative  qualities  are  dominant. 
This  matter  may  be  determined  by  the  time  this  paper  finds  its  way  through 


PLATE   10. 


B 


A.  Raimannia  and  iiuluced  derivatives.     The  three  rows  of  plants  in  the  background  include  some  large 

normal  individuals  and  small  mutants  grown  from  seed  taken  from  a  capsule  which  had  been  treated 
with  calcium  nitrate.     The  remainder  are  mutants  of  the  second  generation. 

B.  Individuals  grown  from  seeds  taken  horn  one  capsule.     The  large  rosette  is  of  the   parental   type,   the 

smaller  are  derivatives,  two  of  which  have  come  into  bloom  at  the  same  age  and  are  scarcely  so  high 
as  the  normal  rosette. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS.      63 

the  press.  Among  the  large  progeny  grown  from  guarded  seeds  was  only  one 
which  showed  any  marked  variation  from  the  blunt  rounded  leaves  of  the 
rosette  of  the  derivative,  but  this  variation  did  not  extend  to  the  foliar  organs 
developed  later. 

The  radium  pencils  which  were  later  used  on  0.  lamarckiana  were  affixed  to 
an  inflorescence  in  such  a  manner  that  the  radium  coatings  were  15  to  25  mm. 
from  developing  flower-buds.  The  corollas  of  many  were  so  retarded  that 
they  failed  to  open  and  fell  off  prematurely.  At  greater  distances  develop- 
ment of  the  ovary  proceeded  but  slowly  and  normal  size  was  not  reached. 
Perfect  seeds  were  formed  in  many  of  them,  however,  and  these  when  sown 
gave  the  normal  frequency  of  the  aberrant  mentioned  above,  and  which  is  not 
to  be  confused  with  the  one  induced  by  the  stimulative  action  of  zinc  sulphate. 

Some  decisive  results  were  also  obtained  from  Raimannia  odorata,  a  member 
of  a  separate  genus  of  the  evening-primrose  family  from  Patagonia.  During 
the  first  season  (1905)  injections  of  the  ovaries  were  made  with  several  sub- 
stances, with  the  result  that  an  atypic  form  identical  in  all  cases  was  found  in 
seeds  from  ovules  that  had  been  treated  in  various  ways.  Two  such  mutants 
were  secured  from  seeds  of  an  ovary  that  had  been  treated  with  a  10  per  cent 
sugar  solution,  10  from  one  that  had  been  injected  with  a  solution  of  calcium 
nitrate  i  part  to  1000  of  distilled  water,  and  one  was  also  found  in  the 
progeny  from  seeds  taken  from  a  capsule  which  had  been  exposed  to  the 
action  of  a  radium  pencil. 

In  all  of  these  injections  ordinary  jointed  metal  syringes  were  used,  and  the 
water  was  from  a  single  distillation  in  a  copper  still,  so  that  no  special  reliance 
may  be  placed  upon  the  purity  of  the  solution  injected  into  the  ovary. 

Seeds  from  capsules  which  had  been  treated  were  harvested  in  June,  1905, 
and  were  sown  early  in  July.  The  atypic  derivatives  could  be  recognized  as 
soon  as  the  cotyledons  were  fully  expanded,  and  no  skill  was  needed  for  their 
detection.  The  parental  form  bore  leaves  villous  hairy,  or  with  ciliate  mar- 
gins, while  the  mutant  was  entirely  and  absolutely  glabrous.  The  leaves  of 
the  parent  generally  have  an  unbalanced  linear  growth  of  the  margins  by 
which  the  blades  become  fluted.  The  excess  of  growth  in  the  mutant  lies 
along  the  midrib  and  the  margins  become  re  volute.  The  leaves  of  the  mutant 
show  much  less  expansion  than  those  of  the  parent,  the  lower  ones  being 
narrowly  linear,  5  to  6  cm.  long  and  3  to  4  mm.  wide,  while  the  upper  ones  are 
lanceolate  linear,  3  to  4  cm.  long  or  even  longer,  the  mutant  being  much  the 
narrower.  The  parental  type  is  of  a  marked  biennial  habit  and  near  the  close 
of  the  season  the  internodes  formed  are  extremely  short,  which  results  in  a 
dense  rosette.  The  mutant  does  not  make  a  rosette,  by  reason  of  the  fact 
that  its  stem  does  not  alter  its  rate  of  elongation,  but  proceeds  at  a  uniformly 
rapid  rate,  thus  presenting  a  leaf-stem  which  appears  to  be  perennial  in  the 
climate  in  which  the  experiments  were  performed.     So  rapidly  did  develop- 


64     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

ment  proceed  that  one  of  the  atypic  individuals  opened  a  flower-bud  on  No- 
vember 27,  1905,  and  seeds  were  harvested  before  the  beginning  of  the  year. 
Other  lots  of  seeds  were  obtained  throughout  the  year  and  several  sowings 
were  made,  with  the  result  that  the  atypic  form  was  found  to  transmit  its  con- 
stellation of  characters  to  its  offspring.  Among  the  fully  atypic  individuals, 
of  which  a  few  hundred  have  been  grown,  one  was  found  with  wide  leaves  like 
the  parental  form,  another  with  ciliate  leaves,  and  another  with  undulate 
leaves.  This  activation  of  parental  characters  is  one  fully  illustrated  by 
lamarckiana  and  its  derivatives  and  shows  that  the  treatment  has  simply 
thrown  certain  parental  characters  into  a  state  of  latency  and  awakened 
others  with  which  the  parental  characters  are  mutually  exclusive  as  to  external 
manifestation.  These  individuals  did  not  afford  a  transition  or  intergrading 
series  between  the  derivative  and  the  parental  form,  however  (plate  10). 

In  the  summer  of  1906  a  second  series  of  injections  was  made,  with  the  result 
that  capsules  treated  with  zinc  sulphate,  i  part  to  2000  parts  of  distilled 
water,  yielded  seeds  which  produced  progeny  inclusive  of  the  atypic  form 
described  above,  and  also  some  other  combinations  which  it  has  not  been 
possible  to  follow  in  successive  generations.  The  group,  however,  bears  a 
general  resemblance  in  relationship  to  that  of  the  lamarckiana  mutants. 

An  injection  with  calcium  nitrate,  i  part  to  1000  of  distilled  water,  was 
without  effect  and  the  progeny  were  all  of  the  parental  type.  It  was  remark- 
able, however,  that  a  treatment  with  this  substance  in  the  previous  season 
had  secured  some  atypic  forms,  probably  due  to  some  opportune  condition  in 
the  experiment  not  yet  understood.  Furthermore,  the  plants  of  the  progeny 
of  the  first  treatment  which  were  apparently  normal  yielded  seeds  which  gave 
a  few  atypic  forms,  indicating  that  the  effect  of  the  first  treatment  had  been 
more  or  less  permanent.  Confirmation  of  this  important  matter  has  not  yet 
been  obtained,  however.  Then,  again,  an  injection  of  ovaries  in  1906  with  the 
distilled  water  subject  to  impurity  from  the  still  and  from  the  syringe,  as  men- 
tioned above,  also  resulted  in  a  progen}^  in  which  some  atypic  forms* were 
found  to  occur.  So  far,  then,  as  experience  with  this  plant  is  afforded  it  is  to 
be  seen  that  a  variety  of  agents  act  in  inducing  discontinuous  variation  in  the 
progeny  and  that  in  one  instance  the  variability  was  carried  to  the  third 
generation,  as  far  as  that  part  of  the  test  has  been  extended.  The  atypic 
forms  transmit  their  qualities  perfectly  from  generation  to  generation,  and 
the  third  generation  now  in  hand  are  like  the  first  from  which  they  came 
originally. 


PLATE  11  . 


:-^    '-^ 


?s... 


B 

A.     Ounotlicra  i^randirtoni,  seedling  alioul  two  months  after  yenninatioii. 
K.      (Jenotlicra  grandiriora,  ro.sette  five  months  ohl. 


IDENTITY   OF   EVENING-PRIMROSES. 

Prepared  by  Anna  Murray  Vail. 

In  view  of  the  extent  to  which  the  evening-primroses  are  being  used  to 
obtain  experimental  evidence  upon  questions  of  heredity  at  the  present  time, 
it  has  been  deemed  important  to  make  a  systematic  study  of  the  group.  To 
this  end  seeds  have  been  obtained  from  correspondents  in  various  parts  of 
America  and  from  European  botanical  gardens  in  which  these  plants  appear 
to  be  extensively  cultivated. 

In  the  few  years  in  which  the  cultures  have  been  under  way  it  has  become 
plainly  apparent  that  a  much  larger  number  of  species  are  native  to  America 
than  has  been  supposed  hitherto.  The  series  of  pedigree-cultures  made  from 
them,  using  carefully  guarded  seeds,  shows  that  botanists  have  customarily 
grouped  many  elementary  species  under  the  name  of  O.  biennis.  Furthermore, 
as  has  been  noted  on  page  8,  none  of  these  appear  to  coincide  with  the  form 
cultivated  in  Europe  under  this  name. 

Several  of  these  elementary  species  occur  on  Long  Island,  a  number  of  them 
collected  by  Mr.  E.  P.  Bicknell  deserving  mention,  and  requiring  special  study 
to  determine  their  relationship  to  the  common  0.  biennis. 

Another  group  of  closely  related  but  apparently  distinct  elementary  species 
came  from  the  west  shore  of  Lake  Champlain.  There,  in  waste  ground  at  the 
roadsides  about  Plattsburg,  fairly  typical  0.  biennis  is  to  be  found.  A  small- 
flowered  and  very  characteristic  plant  abounds  on  inland  sand-dunes,  in 
sandy  clearings  along  the  Ausable  River,  and  on  the  railroad  embankments 
near  Bluff  Point.  In  the  Ausable  River  woods  it  grows  associated  with  0. 
cruciata,  and  a  supposed  hybrid  between  the  two  was  observed.  Occasionally 
a  larger-flowered  form  on  the  edge  of  woods  was  observed  and  still  another  on 
the  gravelly  shores  of  the  lake.  These  types,  with  the  exception  of  the  hybrid 
of  which  no  mature  seed  was  secured,  were  grown  in  the  New  York  Botanical 
Garden,  where  they  reproduced  in  every  particular  the  characters  that  distin- 
guished them  in  the  field.  They  will  require  more  extended  study  for  the 
determination  of  their  specific  relationships. 

The  more  important  facts  concerning  the  anatomy  and  distribution  of  a 
few  forms  have  been  obtained  and  descriptions  of  O.  grandiflora,  0.  simsiana, 
O.  oakesiana,  0.  parviflora,  and  O.  muricata  are  here  presented.  It  is  proposed 
to  reserve  discussion  of  the  large  number  of  other  distinct  forms  until  their 
life-history  and  hereditary  qualities  shall  have  been  more  definitely  ascer- 
tained. 

65 


66     MUTATIONS,    VARIATIONS,    AND  REIvATlONSHIPS   OF   THE   OENOTHERAS. 

The  use  here  of  the  name  Oenothera  for  species  belonging  to  the  genus 
Onagra  (Tournefort,  Adanson,  Spach)  is  according  to  the  decision  of  Dr.  J.  N. 
Rose  in  a  recent  paper  (1905),  where  he  points  out,  and  with  good  reason,  that 
Oenothera  biennis  should  be  considered  as  the  type  of  the  genus.  The  descrip- 
tions in  each  case  have  been  made  from  living  plants. 

OENOTHERA  GRANDIFLORA  Aiton 

Oenothera  grandiflora  Ait.     Hort.  Kew.,  3:   2,  1789. 

Onosiiris  aciiminita  Raf.      Fl.  Ind.,  96,  1817  (?). 

Oenotliera  grandiflora  P  Sims.     Curtis's  Bot.  Mag.  46:   pi.  2068,  1819. 

Oenothera  biennis  var.  grandiflora  Lindl.     Bot.  Regist.  19:  pi.  1604,  1833  (?). 

Onagra  grandiflora  (Ait.)  Vail.     Torreya,  5:   9,  1905. 

Onagra  vulgaris  Spach.     Nouv.  ann.  mus.  Paris,  4:  353,  1835.      (Reprint  33,  1835.) 

In  part. 

Seedling  about  2  months  old. — Rosette  loose,  spreading,  7  to  12  cm.  in  diameter;  outer 
leaves  4  to  7  cm.  long,  2  to  3  cm.  wide;  blades  oblong,  or  oblong-spatulate,  nearly  glabrous, 
irregularly  blotched  with  dull  red,  broader  above  the  middle,  rounding  or  acutish  and 
shallowly  undulate  toothed  at  the  apex,  tapering  at  the  more  deeply  toothed  base  into  the 
margined  petiole ;  the  inner  blades  oblong,  acutish,  and  soon  becoming  more  deeply  toothed 
at  the  base  (plate  11,  a)  . 

Seedling  about  5  vionths  old. — Rosette  rather  loose,  spreading  flat  on  the  ground,  about 
30  cm.  in  diameter;  outer  leaves  12  to  15  cm.  long,  3  to  4  cm.  wide;  blades  oblong-spatulate, 
broadest  above  the  middle,  acutish  at  the  apex,  tapering  at  the  irregularly  pinnatifid- 
toothed  base  into  a  long,  broad  petiole,  obscurely  puberulent  near  the  outer  margin  above 
and  beneath  on  the  white  midvein  and  veins, bright  dark  blue-green,  irregularly  blotched 
throughout  with  red,  which  color  also  appears  sometimes  on  the  wide  white  midvein 
(plate  II,  b). 

Adult  plant. — Plant  1.5  to  3  m.  in  height,  branching  from  the  base  upward,  the  terminal 
portion  of  the  main  stem  bearing  a  cluster  of  short  branches  with  a  profusion  of  flowers. 
Stems  stout,  often  reaching  a  diameter  of  6  cm.  at  base,  nearly  terete  below,  slightly  angled 
and  channeled  above,  or  sulcate,  the  epidermal  tissues  becoming  detached  on  the  lower 
part  of  the  main  stem,  which  is  hard  and  woody,  clothed  in  the  upper  portion  with  short, 
spreading,  somewhat  varyinglymuricate  pubescence;  leaves  10  to  20  cm.  long;  blades  ovate- 
lanceolate,  glabrate,  with  appressed  scattered  hairs  on  the  veins,  denticulate,  being  more 
deeply  and  irregularly  toothed  in  the  basal  portion,  tapering  to  a  short  margined  petiole, 
bright  green  and  shining  above,  paler  beneath;  terminal  rosette-like  cluster  of  the  inflores- 
cence symmetrical ;  flowers  very  abundant  and  fragrant ;  bracts  lanceolate,  3  to  5  cm.  long, 
acuminate,  abruptly  tapering  into  a  short  petiole,  or  sessile ;  conic  portion  of  bud  3  to  4  cm .  long, 
6  to  7  mm.  in  diameter  at  base,  very  slender,  tapering  from  base  to  apex,  sparingly  pubes- 
cent, thin,  dotted  with  small  red  spots,  the  erect  free  tips  very  slender,  setaceous,  8  to  10  mm. 
long;  hypanthium  4.5  to  5.5  cm.  long,  very  slender,  much  longer  than  the  reflexed  calyx- 
lobes  that  usually  cohere  in  pairs  at  the  tips;  ovary  i  cm.  long,  or  less,  slender,  petals  firm, 
3.5  to  4  cm.  long,  and  about  4  cm.  wide,  more  or  less  emarginate,  wedge-shaped  at  base; 
filaments  2.5  cm  long,  very  slender;  anthers  slender,  i  cm.  long;  pistil  much  longer  than 
stamens,  and  projecting  from  the  flower  and  from  the  unopened  buds  late  in  the  season; 
stigmatic  lobes  10  to  12  mm.  long;  capsule  3  to  3.5  cm.  long,  8  mm.  in  diameter  in  thickest 
portion,  green  and  shining,  glabrous  or  with  a  few  scattered  hairs,  four-angled  and  tapering 
from  base  (plate  12). 

Alabama. — Earle's  Landing  and  Dixie  Landing  near  Tensaw,  Tracy,  No.  8001  in  the 
herbarium  of  the  New  York  Botanical  Garden. 


PLATE  12. 


Oenothera  grandiflora.      1,  stem-leaf;  2,  leaf  from  young  rosette;  3,  bract;  4,  bud;  5,  flower 
with  petals  removed;  6,  capsule  and  bract;  7,  petal  (minimum  size). 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS.      67 

Texas. — Wright,  without  locality,  in  the  Gray  Herbarium  of  Harvard  University  (?). 

Kentucky. — Neighborhood  of  Lexington,  Short,  without  date,  in  the  herbaria  of  Columbia 
University  and  Academy  of  Natural  Sciences,  Philadelphia.  The  sheet  in  the  Columbia 
University  herbarium  bears  the  note  "Oenothera  muricata.  Vespertine,  3  to  5  feet  high." 
This  is  the  sheet  referred  to  (MacDougal,  Vail,  Shull  &  Small,  1905,  p.  6)  as  belonging  to 
O.  lamarckiana,  but  which  in  the  writer's  opinion  should  belong  here.  The  Philadelphia 
specimen  gives  the  exact  locality  "Neighborhood  of  Lex"  fl.  Augt. &  after,  i  mile  from  town 
left  of  Coles  road  to  Frankfort. "     It  may  be  an  escape  from  cultivation. 

The  rediscovery  of  the  original  type  locaHty  of  this  species  is  recorded  in  a 
previous  paper  (MacDougal,  Vail,  Shull  &  Small,  1905,  pp.  7  and  8).  The 
seeds  from  which  the  plants  described  above  were  raised  were  sent  to  the  New 
York  Botanical  Garden  from  Tensaw,  Alabama,  by  Mrs.  J.  F.  Davis  at  the 
request  of  Professor  Tracy,  absolutely  ripe  capsules  not  having  been  found  at 
the  time  of  his  visit  late  in  August,  1904.  In  cultivation,  0.  grandiflora  has 
the  appearance  described  by  Bartram  in  1793  and  again  by  Professor  Tracy  in 
1904.     (MacDougal,  Vail,  Shull  &  Small.  1905,  pp.  7  and  8). 

It  is  very  fragrant  and  showy,  and  flowers  abundantly.  Most  conspicuous 
and  characteristic  are  the  tapering  pale-yellow  buds  with  long,  slender,  seta- 
ceous tips  to  the  calyx-lobes.  They  have  none  of  the  heavy,  swollen  appear- 
ance of  those  of  O.  lamarckiana  and  of  some  of  its  derivatives.  For  24  hours 
or  so  before  the  opsning  of  the  flower  the  closed  stigma  crowded  into  the  apex 
of  the  bud  is  distinctly  seen  through  the  thin  epidermis  of  the  closed  calyx- 
lobes.  The  calyx-lobes  when  expanded  are  split  open  in  twos  (but  very  rarely 
in  fours),  the  petals  only  attaining  their  full  size  during  expansion.  This  is 
also  the  case  in  Oenothera  argillicola  (Onagra  argillicola  McKenzie).  (Mac- 
Dougal, Vail,  Shull  &  Small,  1905,  p.  12.) 

A  further  comparison  of  supposed  plates  of  0.  grandiflora  with  living  speci- 
mens of  the  species  would  indicate  the  following :  The  plate  of  ' '  Oenothera 
lamarckiana"  in  Ivcmaire  (Illustration  horticole,  9,  p.  318,  1862)  appears  to 
be  that  species,  but  the  description  of  the  600  flowers,  buds,  and  capsules  on 
one  single  plant  would  appear  to  refer  to  0.  grandiflora,  2iS  O.  lamarckiana  does 
not  (in  cultivation)  flower  so  abundantly.  The  plate  in  Edward's  Botanical 
Register  No.  1604  of  ''Oenothera  biennis  var.  grandiflora"  has  much  stouter 
bud-tips,  bracts  that  are  broader  at  the  base,  and  much  broader  petals  than 
the  typical  O.  grandiflora. 

The  plate  Oenothera  grandiflora  Sims  (Curtis's  Bot.  Mag.,  pi.  2068)  is  quite 
typical  of  the  plant  grown  in  the  New  York  Botanical  Garden  as  to  the  shape 
of  the  petals,  but  the  bud-tips  are  much  too  heavy  and  the  bracts  too  broad 
and  clasping  at  the  base. 

Large-flowered  evening-primroses  have  appeared  from  time  to  time  in  the 
eastern  seaboard  States;  but  none  that  have  been  examined  so  far  can,  be 
determined  as  being  certainly  indigenous  there.  In  the  Gray  Herbarium  the 
following  specimens  are  noted. 


68     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS    OF   THE   OENOTHERAS. 


New  Jersey. — Pine  barrens,  Miss  Treat  of  Vineland,  1871,  "wild."  This  has  a  flower 
that  when  expanded  must  have  measured  nearly  10  cm.  in  diameter,  the  petals  being  5  cm. 
long  and  over,  but  the  bud  is  stouter  and  heavier  than  0.  grandiflora  and  has  more  the  char- 
acter of  that  of  O.  lamarckiana.  It  might  be  interesting  to  know  whether  it  still  survives  in 
the  pine  barrens. 

Maine. — Orono.  M.  L.  Fernald,  July  17,  1892.  A  specimen  with  very  much  the  appear- 
ance of  Oenothera  rubrinervis,  having  the  very  characteristic  bud  of  that  derivative  of  O. 
lamarckiana.  It  is  probably  not  indigenous.  Large-flowered  specimens  labeled  Oenothera 
biennis  var.  grandiflora  and  Oenothera  grandiflora  from  California  appear  to  be  Oenothera 
hookeri  Torrey  &  Gray. 

OENOTHERA  SIMSIANA  SeringE. 

Oenothera  corymbosa  Sims.  Curtis's  Bot.  Mag.,  45:  pi.  1974,  1818.  Not  Lamarck. 
Oiiagra  spectabilis  Spach.  Nouv.  ann.  mus.  Paris,  4:  352,  1835.  (Repr.  32,  1836.) 
Oenothera  simsiana  Seringe,  in  DC.  Prodr.,  3:  47,  1828. 

Seedling  about  6  weeks  old. — Leaves  glabrate;  blades  oblong,  those  of  the  later  leaves  oval, 
20  to  35  mm.  long,  10  to  15  mm.  wide,  obtuse  at  the  apex,  gradually  tapering  into  the 
margined  petiole,  light  yellow-green  (fig.  72). 

Seedling  about  5  months 
old. — Rosettes  rather  loose, 
40  to  45  cm.  in  diameter, 
spreading;  outer  leaves  15  to 
20  cm.  long,  4  to  6  cm.  wide; 
blades  oblong  to  obovate, 
broadest  somewhat  above  the 
middle,  taperingto  the  obtuse 
apex  or  in  the  younger  leaves 
to  an  acutish  apex,  gradu- 
ally tapering  into  the  broad 
petiole,  light  yellow-green, 
obscurely  puberulent,  with 
widely  scattered  hairs  on 
both  surfaces  (under  a  lens), 
approximately  repand- 
denticulate  with  rather  dis- 
tinct shallow  teeth;  petiole 
white,  8  mm.  or  more  wide. 

Mature  rosette. — Leaves  loosely  spreading,  finely  but  obscurely  pubescent  all  over,  the 
larger  ones  20  to  28  cm.  long,  5  to  6  cm.  or  more  wide;  blades  oblong  to  oblong-obovate, 
shallowly  repand-denticulate,  tapering  into  a  very  wide  white-margined  petiole  (plate  13). 

Adult  plant. — Plant  rather  straggling  in  appearance,  appressed-pubescent  and  hispidu- 
lous,  especially  in  the  upper  portions.  Main  stem  about  2  m.  in  height,  greenish-white, 
becoming  reddish,  stout,  terete,  or  slightly  angled  and  sulcate  above,  irregularly  branched 
at  the  base,  the  long,  rather  weak,  virgate  branches  ascending;  leaves  minutely  pubescent 
above,  somewhat  less  so  beneath,  7  to  10  cm.  long,  blades  obscurely  repand-denticulate, 
oblong-lanceolate  to  lanceolate,  acute  or  acuminate  at  the  apex,  abruptly  narrowed  into  a 
very  short  margined  petiole  or  the  uppermost  sessile,  coriaceous,  very  brittle,  bright  yellow- 
green,  soon  turning  red,  twisted,  the  upper  ones  more  markedly  so;  terminal  rosette-like 
cluster  of  the  inflorescence  symmetrical,  the  floral  bracts  divaricate  and  somewhat  reflexed; 
bracts  lanceolate,  acuminate,  subsessile  and  subcordate;  conic  portions  of  the  bud  2  cm. 
long,  6  to  7  mm.  in  diameter  at  the  base,  canescently  appressed-pubescent  and  hirsute, 


KiG.   72. — Oenothera  simsianasecdling  about  6  weeks  after  germination. 


PLATE   13. 


Oenothera  simsiana.      1,  leaf  from  mature  rosette;    2,  stem-leaf;   3,  bud;  4,  flower  with 
petals  removed  and  bract;  5,  capsule  and  bract;  6,  petal. 


PLATE  14. 


t.\ 


■■:;< 


Adult  rosctti.:  of"  Oenothera  simsiana    (upper  left-hand  corner).      Mature  plant  of  Oenothera  sinisiana  in 
experimental  s^rounds  in  New  York,  hearing  flowers  and  capsules. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE   OENOTHERAS.     69 

tapering  in  terminal  portion  only,  the  erect,  free  tips  of  the  sepals,  3  to  4  mm.  long,  mostly- 
unequal;  hypanthium  4  to  5  cm.  long,  very  slender,  densely  appressed-pubescent;  ovary 
1 2  to  13  mm.  long,  slender,  densely  appressed-pubescent ;  sepals  3  to  4  cm.  long,  shorter  than 
the  tubular  portion  of  the  hypanthium ;  petals  thin,  bright  golden-yellow,  fading  saffron- 
yellow,  4  cm.  long,  4  to  5  cm.  wide,  deeply  emarginate;  filaments  25  mm.  long,  very  slender; 
anthers  slender,  12  to  15  mm.  long;  pistil  as  long  or  slightly  longer  than  the  stamens;  lobes 
of  the  stigma  6  to  7  mm.  long,  divaricate;  capsule  about  3  cm.  long,  6  to  7  mm.  in  diameter 
at  the  widest  portion,  4-angled,  appressed-pubescent,  rather  abruptly  contracted  at  the 
apex  (plate  14). 

Type  locality  of  the  plants  described  above,  along  International  Railroad, 
City  of  Mexico.  Collected  by  J.  N.  Rose  and  Jos.  H.  Painter,  September  20, 
1903.     No.  7219. 

Seeds  of  a  late-flowering  plant  were  sent  to  the  New  York  Botanical  Garden, 
unnamed,  by  Dr.  Rose  late  in  1904,  and  the  plants  from  which  the  above 
description  was  compiled  were  grown  under  glass  for  the  first  6  months  and 
then  transferred  to  the  experimental  grounds.  The  flowers  produced  at  the 
height  of  maturity  are  somewhat  larger  than  those  received  with  the  seeds, 
but  the  late  summer  flowers  exactly  resembled  those  from  the  original  locality. 
Remarkable  for  the  form  and  color  of  the  light  green  rosette  and  for  the  very 
characteristic  assurgent  or  erect  habit  of  the  slender  virgate  stems  and  the 
acuminate,  curved,  and  twisted  leaves. 

This  species  is  referred  to  Oenothera  simsiana,  perhaps  somewhat  doubt- 
fully. The  plate  (1974)  in  Botanical  Magazine  agrees  with  it,  except  for  the 
corymbose  inflorescence,  which  may  occur  readily  enough  in  the  plant  in  its 
native  habitat,  as  many  related  species  var}"  in  that  regard.  The  mature  and 
normal  capsules  do  not  show  the  strongly  reflected,  white  valve-tips  that  are 
seen  in  the  illustration,  but  the  plant  as  cultivated  in  the  New  York  Botanical 
Garden  was  subject  to  the  sting  of  an  insect  that  caused  a  malformation  of  the 
ovary  or  young  capsule,  which  then  had  very  much  the  appearance  of  the  cap- 
sules in  Sims's  plate,  though  when  malformed  they  never  quite  reached  the 
state  of  maturity  depicted  there.  This  is  what  may  have  occurred  at  Long- 
leats  in  18 16. 

0.  simsiana,  with  the  synonymy  as  given  above,  is  listed  in  Hemsley  (Biol. 
Centr.  Am.,  1 :  454)  without  any  indication  of  definite  locality  or  collection. 

An  unnamed  Oenothera  collected  at  the  city  of  Durango  and  vicinity,  by 
Dr.  Edward  Palmer  (No.  293,  1896,  in  the  herbarium  of  the  New  York  Botani- 
cal Garden)  can  be  referred  here  with  a  fair  degree  of  certainty. 

O.  simsiana  is  closely  related  to  Oenothera  hookeri  Torrey  &  Gray,  with 
which  it  has  doubtless  been  confounded  in  herbaria.  The  growing  plants  are, 
however,  very  dissimilar  from  that  species  in  general  aspect  and  habit  from 
the  rosette  stage  to  maturity,  and  the  flower  appears  to  retain  its  yellow  color 
very  much  longer  than  does  0.  hookeri. 


70     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


OENOTHERA  OAKESIANA  (Robbins)  S.  Watson. 

Oenothera  biennis  var.  oakesiana  Robbins.     A.  Gray,  Man.,  ed.  5,  178,  1867. 
Oenothera  oakesiana  S.  Watson.     Biblio.  Ind.  N.  Am.  bot.,  383,  1878. 
Onagra  oakesiana  Britton.     Mem.  Tor.  bot.  cl.,  5:  233.  1894. 

Seedling  about  2  months  old. — Leaves  obscurely  puberulent,  with  few  scattered  hairs; 
blades  oblong  or  oblong-obovate,  6  to  8  mm.  v.ide,  rounded  or  obtuse  at  the  apex,  tapering 
to  a  slender  margined  petiole  (plate  15,  a). 

Seedling  about  5  months  old. — Rosette  crowded,  symmetrical,  7 to  8  cm.  in  diameter;  outer 
leaves  3  to  4  cm.  long,  8  to  10  mm.  wide;  blades  oblong-ovate,  broadest  above  the  middle, 
obtuse  at  the  apex,  tapering  to  the  white,  margined,  entire  petiole,  dull  or  pale  blue-green, 
rather  fleshy,  approximately  denticulate  above,  glabrous,  except  for  a  few  obscure  hairs  on 
the  margins. 

Seedling  8  months  old. — Rosette  9  to  10  cm.  in  diameter,  very  symmetrical  and  crowded, 
raised  above  the  ground  with  a  few  of  the  old  leaves  adhering  to  the  rootstock;  blades  approx- 
imately and  more  prominently  denticulate. 

Mature  rosette. — Leaves  more  or  less  strigose-pubescent  all  over,  the  central  ones  quite 
densely  so,  the  larger  ones  from  15  to  20  cm.  long;  blades  narrowly  lanceolate,  approxi- 
matelyand  shallowly  toothed  at  the  acute  apex,  more  deeply  toothed  at  the  slender,  taper- 
ing base  (plate  15,  b). 

Adult  plant. — Plant  i  to  1.5  m.  high,  rather  slender,  clothed  nearly  throughout  with  a  fine, 
close,  strigose,  appressed,  and  somewhat  cinereous  pubescence  and  few  scattering  longer 
spreading  hairs,  becoming  almost  glabrate  and  glaucous  with  age.  Stem  light  or  whitish 
in  color,  angled  and  channeled  above,  branched  to  the  middle;  leaves  10  to  15  cm.  long; 
blades  shallowly  and  remotely  toothed,  often  more  deeply  so  at  base,  narrowly  lanceolate, 
acute  and  tapering  at  each  end,  sessile  or  nearly  so,  gray-green  and  shining  above,  paler 
beneath,  rather  thick  and  somewhat  brittle;  terminal  rosette-like  cluster  of  the  inflorescence 
symmetrical,  the  floral  bracts  very  small,  divaricately  spreading;  bracts  narrowly  lanceolate, 
more  deeply  and  regularly  toothed  than  the  leaves,  acuminate  at  the  apex,  tapering  to  the 
base,  sessile,  becoming  2.5  times  as  long  as  the  mature  capsule;  conic  portion  of  bud  10  to 
13  mm.  long,  rather  prominently  4-angled,  4  to  5  mm.  in  diameter  at  the  base,  the  free 
separate  divaricate  tips5mm.  long,  finely  but  inconspicuously  appressed  pubescent;  hypan- 
thium  2.5  mm.  to  2.7  mm.  long,  slender,  finely  but  sparingly  pubescent;  calyx-lobes  half  as 
long  as  hypanthium;  petals  firm,  13  to  15  mm.  long,  12  to  14  mm.  wide,  deeply  emarginate, 
not  opening  widely;  filaments  12  mm.  long;  anthers  7  mm.  long;  pistil  a?  long  or  slightly 
shorter  than  the  exserted  stamens,  the  lobes  erect,  4  to  5  mm.  long;  capsules  3  to  3.5  mm. 
or  more  long,  7  to  9  mm.  in  diameter,  angled  and  rounded,  finely  appressed-pubescent, 
abruptly  constricted  near  the  apex.     Remarkable  for  its  large  seeds  (plates  t6  and  17). 

Massachusetts. — Uxbridge,  J.  \V.  Robbins,  Au.gust,  1878. 

Rhode  Island. — Providence,  E.  P.  Bicknell,  August,  1896;  September,  1899. 

New  York. — New  York  Botanical  Garden,  D.  T.  MacDougal,  September,  1904;  Cold 
Spring  Harbor,  Long  Island,  G.  H.  Shull,  September,  1904. 

The  above  specimens  are  the  only  ones  in  the  herbaria  of  the  New  York 
Botanical  Garden  and  Columbia  University  that  can  with  any  degree  of 
certainty  be  referrred  to  0.  oakesiana.  The  Robbins  specimen  is  from  the 
herbarium  of  the  late  Rev.  Thomas  Morong  and  attached  to  its  sheet  is  the 
following  note  in  the  writing  of  its  discoverer. 


PLATE   15. 


B 


A.  SeL-dliiiij  of  Oenotlifni  (>:ikt;sian:i  about  two  inoiitlis  old 

B.  Ailult  rosette  of  Oenothera  oakesiana. 


PLATE  16. 


Oenothera  oakesiana.      l.leaf    from  mature    rosette;    2,    stem-leaf;    3,  capsule    and    bract; 
4,  flower  with  petals  removed;  5,  buds;  6,  petal  (minimum  size). 


PLATE  17. 


()eii(>tlic'i:i  oake^iaiKi  "rown  wild  in  waste  land  in  New  \'i)rk  Botanical  (iardcn. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE   OENOTHERAS.      7 1 

Oenothera  oakesiana  mihi. 

This  plant  was  presented  to  Professor  Gray  many  years  ago  as  a  well-characterized  and 
distinct  species,  which  I  still  consider  it.  I  discovered  it  in  the  medical  colleges  in  New 
Havea,  Connecticut,  in  1S27.  In  1831  1  sowed  its  seed  with  those  of  the  ordinary  variety. 
The  differences  are  as  follows: 

(i)  If  raised  from  seeds  sown  in  the  spring  it  is  annual,  the  common  variety  by  its 
side  being  biennial. 

(2)  The  leaves  of  the  first  year,  when  biennial,  are  much  narrower. 

(3)  The  pubescence  throughout  is  soft-appressed,  that  of  the  other  being  coarser  and 
spreading. 

(4)  The  points  of  the  calyx  are  always  spreading,  in  the  other  appressed. 

(5)  The  capsule  is  longer  and  more  taper. 

(6)  The  ripe  seeds  are  larger. 

I  have  found  but  two  other  localities,  namely,  on  sides  of  the  railroad  north  of  Norton, 
Massachusetts,  and  another,  same  situation  near  Apponong  Depot,  Rhode  Island.  Also  it 
is  well  naturalized  in  Uxbridge,  Massachusetts,  from  my  seeds. 

UXBRIDGE,  5th  Sept.,   1878.  J.  W.  ROBBINS. 

The  plants  from  the  New  York  Botanical  Garden  were  first  noticed  by  Dr. 
MacDougal  late  in  the  summer  of  1904.  They  grew  not  far  from  the  embank- 
ment of  the  Harlem  Division  of  the  New  York  Central  Railroad  and  were 
quite  abundant,  though  not  covering  a  very  widespread  area.  With  them 
were  associated  a  number  of  0.  biennis,  the  latter  in  that  particular  spot,  how- 
ever, being  not  as  abundant  as  elsewhere  in  the  garden.  Rosettes  were 
obtained  of  the  O.  oakesiana  and  the  plants  raised  from  them  the  following 
summer  were  in  every  way  similar  to  the  wild  plants,  and  formed  the  basis  of 
these  notes  on  the  species.  Plate  17  was  photographed  from  a  specimen 
growing  wild  in  the  New  York  Botanical  Garden. 

OENOTHERA    PARVIFLORA    Linnaeus. 

Oenotliera  parviflora  Linnaeus.     Syst.,  ed.  10:   998,  1759. 
Onagra  parviflora  Moench.      Meth.  Supp!.,   287,    1802. 

Seedling  about  6  weeks  old. — Rosette  2.5  to  3  cm.  in  diameter,  rather  loose  and  irregular, 
blades  of  the  leaves  ovate  or  ovate-spatulate,  glabrate,  ciliate. 

Rosette  about  3  months  old. — Leaves  finely  pubescent  on  both  surfaces,  less  so  above; 
blades  various,  those  of  the  earlier  leaves  narrowly  oblong  or  oblong-spatulate,  varying  to 
oblong-obovate,  6  to  10  cm.  long,  the  larger  ones  2.5  to  3  cm.  wide,  approximately  and 
shallowly  denticulate,  obtuse  or  nearly  so  at  the  apex,  gradually  narrowed  to  the  base  of  the 
margined  petiole,  which  is  very  early  streaked  with  red  or  pink. 

Mature  rosette. — Leaves  long  and  slender,  spreading  flat  on  the  ground;  blades  lanceolate, 
or  oblong-lanceolate,  acutish,  irregularly  and  strongly  denticulate,  more  deeply  toothed  at 
the  long,  tapering  base,  dark  green  and  shiny,  sparingly  mottled  with  red,  sparingly  puber- 
ulent  above,  more  so  and  paler  beneath,  15  to  30  cm.  long,  3  to  5  cm.  wide;  petiole  white, 
8  mm.  wide,  margined  to  base,  distinctly  pink  or  red  ringed  most  of  its  length  (plates  18 
and  20). 

Adult  plant. — Central  stem  not  always  maturing  the  first  season  when  forced.  Lateral 
branches  of  annual  rosette  stout  and  angled,  pubescent  with  spreading  hairs;  leaves 
resembling  those  of  the  rosette  in  color,  texture,  and  pubescence;  blades  oblong  to  oblong- 


72     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE    OENOTHERAS. 

lanceolate,  tapering  to  the  short,  margined  petiole,  shallowly  and  approximately  denticulate 
above,  some  of  them  crinkled  close  to  the  midvein;  terminal  rosette-like  cluster  of  the  inflo- 
rescence symmetrical  and  spreading;  bracts  lanceolate,  acute,  sessile  or  nearly  so,  divaricate, 
crowded  on  short  internodes;  flowers  numerous;  bud  club-shaped,  the  conic  jjortion  8  mm. 
long,  3  to  4  mm.  in  diameter,  rarely  angled,  the  free  tips  separate,  quite  slender,  erect,  2  nmi. 
long  or  less,  mostly  tipped  with  red:  hypanthium  3  to  3.5  mm.  long,  slender,  i)uberulent; 
ovary  8  to  9  mm.  long;  calyx  lobes  one-third  as  long  as  the  hypanthium;  petals  firm,  S  mm. 
long,  9  mm.  wide,  deeply  emarginate,  cuneate;  filaments  7  mm.  long;  anthers  4  to  5  mm. 
long,  pistil  shorter  than  the  slightly  exserted  stamens,  the  lobes  divaricate,  3  to  4  m.m.  long, 
capsules  abundant,  2.5  cm.  long,  6  mm.  in  diameter  at  the  widest  portion,  bluntly  4-angled, 
pubescent,  bright  green,  much  thinner  in  apical  portion.  (The  apex  of  some  of  the  valves 
are  occasionally  more  or  less  distinctly  bifid.)  (Plate  19.) 
Illustratioti. — Meerburg,  Plantas  rariores,  pi.  34,  1789. 

Early  in  1906  a  package  of  seeds  distributed  as  Oenothera  parviflora  was 
received  from  the  Botanic  Garden  of  Madrid,  Spain.  The  plants  raised  from 
these  seeds  differed  essentially  from  any  of  the  various  so-called  O.  biennis  of 
European  origin  in  cultivation  in  the  experimental  grounds  of  the  New  York 
Botanical  Garden.  The  plants  were  very  persistently  biennial,  and  only  one 
or  two  of  those  placed  in  the  open  threw  out  lateral  branches,  the  main  branch 
only  starting  its  growth  too  late  in  the  autumn  to  mature.  Specimens  of  any 
Oenothera  closely  resembling  it  are  not  to  be  found  in  the  herbarium  of  the 
garden  or  in  that  of  Columbia  University.  As  0.  parviflora  Linnaeus  was 
credited  to  "Canada  to  Virginia"  by  Pursh  (Fl.  Am., Sept.,  261, 1814),  with  the 
further  note  that  it  was  rare,  the  development  of  the  Madrid  plants  was  fol- 
lowed and  recorded  with  the  hope  that  eventually  the  species  might  still  be 
found  a  component  part  of  the  North  American  flora. 

Late  in  the  fall  of  1905,  Miss  N.  M.  Stevens  sent  in  some  rosettes  of  what  she 
considered  as  two  distinct  species  of  Oenothera  collected  in  the  neighborhood  of 
South  Harpswell,  Maine.  The  rosettes  were  placed  in  the  experiment?  1 
greenhouse  for  the  winter  and  later  Miss  Stevens  sent  seeds  of  both  species 
from  the  same  locality.  These  latter  germinated  in  the  greenhouse  and 
though  somewhat  similar  in  the  early  rosette  stage,  one  of  them,  the  "pre- 
vailing species"  of  the  locality  from  whence  the  seeds  came,  was  identified 
with  the  plant  determined  as  0.  muricata  Linnaeus  and  the  other,  the  "rarer 
species"  growing  on  sandy  wastes,  was  identical  with  the  species  grown  from 
the  Madrid  garden  seeds  under  the  name  of  0.  parviflora,  and  it  is  from  the 
plants  raised  from  Miss  Stevens's  seeds  that  this  description  has  been  drawn 
(plate  20). 

The  species  shows  distinct  characteristics  in  color  and  habit.  The  flowers 
are  but  rarely  exserted  beyond  the  bracts,  which  are  so  crowded  on  short 
internodes  that  the  ends  of  the  branches  have  a  closely-tufted  appearance. 

The  Linnaean  description  under  Oenothera  reads:  "Parviflor.  A.  OE.  fol. 
ovate-lanceolatis  planis,  caule  laevi  subvelloso.  Margo  coronans  jructum,  non 
uti  praecedentis  quadrifldus ,  sed  octofidus." 


PLATE  18. 


\.A 


Ocnotlicni  p;n-viriora  gn)\vn  from  seeds  obtained  from  the  notanical  Garden  of  Madrid. 


FL4.TE  19. 


2  1 

Oenothera  paniflora.  1,  seedling  two  months  after  germination;  2,  leaf  of  young  rosette; 
3.  stem-leaf;  4,  bud  and  bract;  5,  flower  with  petals  removed;  6,  flower  and 
bract;  7,  petal;  8,  capsule  and  full-grown  bract  (from  seeds  from  Maine). 


PLATE  20. 


*i^•^-*- 


•f 


(Xiiotlicra   p:ii\  itlora  grown  from  scuils  ot  plants  native  near  Soulli  Ilarpswell,  Maine. 


PLATE  21. 


Oenothera  muricata.  1,  leaf  from  mature  rosette;  2,  stem-leaf;  3  and  4,  young  bracts  and 
buds;  5,  flower  with  petals  of  maximum  size;  6,  petal;  7,  flower  with  petals 
removed;  8,  capsule  with  bract;    9,   capsule. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE)   OENOTHERAS.      7J 

The  last  character  would  appear  to  be  an  accidental  one,  that  is,  if  the  name 
belongs  at  all  to  the  plant  in  question  here,  and  it  is  extremely  doubtful  if 
that  point  can  ever  be  definitely  settled. 

Some  of  the  capsules  of  the  Madrid  plant  showed  a  distinct  bifid  termination 
of  the  valves  of  the  capsules,  but  in  the  Maine  plant  that  character  was  less  dis- 
tinct or  altogether  lacking.  The  Spanish  plants  and  those  from  Maine,  grow- 
ing side  by  side,  were  not  distinguishable,  but  the  Maine  plants  matured  more 
rapidly. 

OENOTHERA  MURICATA  Linn^us. 

Oenothera  muricaia  Linnaeus.     Syst  ,  ed.  12,  26.^,  1767. 
Onagra  muricaia  Moench.      Metli.,  675,  1794. 

Seedling  about  6  weeks  old. — Rosette  crowded  and  a  little  irregular,  2  to  3  cm.  in  diameter, 
blades  of  the  leaves  oblong,  glabrate,  slightly  ciliate  on  the  margins. 

Seedling  about  3  months  old. — Rosettes  crowded,  7  to  8  cm.  in  diameter;  leaves  3  to  3.5  cm. 
long,  7  to  8  mm.  wide,  blades  oblong-spatulate,  broadest  above  the  middle,  obtuse  at  the 
apex,  tapering  into  the  margined  petioles,  blue-green,  fleshy,  glabrate  or  becoming  appressed- 
pubescent  as  the  leaves  increase  in  size,  obscurely  denticulate,  the  apex  of  the  denticulations 
mostly  reddish. 

Mature  rosettes  about  5  months  old. — Rosetts  15  to  17  cm.  in 
diameter,  rootstock  thick,  raised  2  to  3  cm.  above  the  ground, 
bearing  a  few  old  leaves  under  the  later  green  ones;  leaves 
crowded,  the  outer  9  to  14  cm.  long,  1.5  to  2  cm.  wide;  blades 
approximately  denticulate,  irregularly  and  more  deeply  so  towards 
the  base  above  the  broad,  white-margined  petiole,  appressed- 
pubescent  or  hirsute,  v>ithout  any  trace  of  red  in  the  leaves, 
except  at  the  apex  of  the  denticulations. 

Adult  plant. — Plant  1.5  m.  high,  pubescent  with  short  appressed 
as  well  as  longer  spreading  hairs  throughout.     Stems  erect,  stout, 

1  to  1.5  m.  high  or  more,  single  or  generally  branched  above, 
angled,  turning  red  and  the  outer  bark  splitting  into  shreds  at 
maturity;  stem-leaves  rather  crowded,  loto  15cm.  long,  1.5  to 

2  cm.  wide;  blades  narrowly  lanceolate  or  linear-oblong,  acute, 
approximately  denticulate,  more  deeplyso  atthebase,  appressed- 

pubescent  with  short  strigose  hairs  more  or  less  on  each  surface,  ^^^-    73.— Seedling    of    Oeno- 

,.,  .,  ,.,,  J.,  Ihera     muricaia    about     2 

light  green,  paler  beneath;  termmal  rosette-like  cluster  or  the  months  old. 
inflorescence  barely  symmetrical;  midvein  broad,  whitish;  inflo- 
rescence crowded;  bracts  lanceolate,  acute  at  the  apex,  tapering  to  the  sessile  base, 
finally  two  or  more  times  as  long  as  the  mature  capsule;  conic  portion  of  bud  1 1  to  12  mm. 
long,  4  mm.  in  diameter,  pubescent  with  appressed  and  spreading  hairs,  the  free  tips  4  mm. 
long;  hypanthium  2.5  to  3  cm.  long,  pubescent;  calyx-segments  about  half  as  long  as  the 
hypanthium;  ovary  i  cm.  long;  petals  thick,  10  to  12  mm.  long,  11  to  14  mm.  wide,  deeply 
emarginate,  not  opening  widely;  filaments  about  i  cm.  long;  anthers  4  to  5  mm.  long;  pistil 
shorter  than  stamens;  stigmatic  lobes  erect,  thickish,  3  to  4  mm.  long  (plate  21). 

This  description  is  based  on  plants  raised  from  seeds  secured  from  Long 
Island  (Bicknell,  1904,  and  Vail,  1904)  and  grown  two  successive  years  in  the 
New  York  Botanical  Garden. 


74     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF    THE    OENOTHERAS. 

Tha  Linnsean  description  reads  as  follows:  "OE.  fol.  lanceolatis  planis, 
caule  purpurascente  muricato.  Similis  parviflorae,  sed  Fructus  os  non  S-fidis. 
Caulis  puncta  rubra  sparsa.     Canada."     (L.  Syst.,  ed.  12,  263.      1767.) 

Based  on  this  description  and  without  reference  to  any  later  interpretation 
of  the  species,  the  following  specimens  are  placed  here : 

Maine. — York,  Bicknell,  1896;  Cape  Neddick,  Bickiiell,  1896;  South  Harpswell,  Stevens, 
1905. 

Massachusetts. — Ipswich,  Morong,  1872;  Nantucket,  Bicknell,  1899. 

New  York. — Long  Island,  Edgemere,  Bicknell,  1902  and  1904;  Coney  Island,  Vail,  1904; 
Staten  Island,  New  Dorp,  Kearney,  1894. 

0.  muricata  L.  raised  from  seed  received  from  Professor  De  Vries  from  the 
Holland  sand-dunes  resembled  these  American  plants,  but  were  not  abso- 
lutelv  identical.  This  should  also  be  said  for  some  plants  that  were  raised 
from  seed  collected  by  Professor  De  Vries  near  Chicago,  Illinois,  in  1904. 
Undoubtedly  0.  muricata  has  a  wider  distribution  in  the  northeastern  States 
than  can  be  noted  here. 

In  addition  to  these  quite  a  number  of  plants  that  apparently  are  referable 
to  this  species  are  preserved  in  the  herbaria  of  the  New  York  Botanical 
Garden  and  of  Columbia  University.     They  are  as  follows: 

Canada. — Province  of  Quebec,  New  Carlisle,  Williams  &  Fernald,  1902;  Rimouski 
County,  Collins  &  Fernald,  1904;  River  Ste.  Anne  des  Monts,  Collins  &  Fernald,  1905; 
Anticosti,  Jupiter  River,  Macoun,  1 883. 

Massachusetts. — Provincetown,  Hollick,  1901;  Nantucket,  MacDougal,  1905. 

Rhode  Island. — Block  Island,  Hollick,  1897. 

The  Nantucket,  Provincetown,  and  Block  Island  plants  are  probably 
stunted  specimens  merely,  but  the  Canada  specimens  are  quite  remarkable 
in  that  the  mature  plant  still  preserved  the  remains  of  the  rosette  of  the  pre- 
vious year  at  the  base  of  the  stem  and  in  general  appearance ;  even  to  the  bent 
habit  of  the  upper  portion  of  the  stem  they  can  be  easily  identified  with  the 
plate  and  description  of  0.  muricata  in  Flora  Danica,  pi.  1757,  1823. 

An  attempt  was  made  to  raise  some  plants  from  seed  taken  from  Messrs. 
Collins  &  Fernald's  specimen  from  River  Ste.  Anne  des  Monts,  1905,  but  it 
was  not  very  successful.  The  plants  were  quite  persistently  biennial  and  in  no 
case  was  a  normal  central  stem  secured,  and  the  rosettes  in  the  garden  were 
very  much  larger  than  those  on  the  herbarium  specimens.  The  Canada  plants 
are  evidentlv  a  more  northern  form  of  the  Long  Island  plant  (plate  22). 

The  habit  of  the  rosette  of  the  previous  year  remaining  at  the  base  of  the 
flowering  stem  of  the  second  year's  growth  is  one  that  is  also  claimed  for  his 
species  0.  ammophila  by  O.  Focke  (1904). 

The  plate  of  0.  muricata  published  by  Murray  (Nov.  comm.  soc.  reg.  Sci. 
Goett.,  6:  24,  pi.  i,  1776)  appears  to  have  broader  leaves  than  the  American 
plants  enumerated  here. 


PLATE  22. 


Ounolbcra  inuiicata  grown  iR-;ir  the-  nortlicrn  limits  of  the  species  in  (^lebcc. 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS.      75 


GENERAL  DISCUSSION. 

HISTORICAL  CONSIDERATIONS. 

With  the  increasing  attention  being  enhsted  in  the  subject  of  mutations  in 
organisms,  numerous  records  of  the  saltatory  activation  and  latency  of  char- 
acters in  lines  of  descent  are  being  disclosed  by  workers  in  all  branches  of  bio- 
logical science.  Some  observations  published  by  Dr.  Arthur  Hollick  in  March, 
1879  (Holhck,  1879)  are  among  the  most  suggestive.  The  views  expressed  by 
Dr.  Hollick  over  26  years  ago  harmonize  so  well  with  the  sum  of  informa- 
tion available  at  the  present  time  that  it  will  be  profitable  to  give  them  in 
full  in  this  place.      He  saj^s: 

The  object  is  more  to  call  attention  to  a  few  facts  which  seem  to  have  been  generally 
passed  over  in  botanical  researches  as  devoid  of  interest.  The  whole  subject  arranges 
itself  under  two  heads.  The  first  will  comprise  true  "albinos"  or  such  plants  as  have 
spontaneously,  in  a  state  of  nature,  lost  their  colors  and  become  white  flowered.  The  second 
relates  to  those  plants  in  which  the  colors  have  been  more  or  less  eliminated  by  artificial 
means. 

First,  then,  we  have  to  consider  those  "sports"  of  nature  where  there  has  been  a  sudden 
change,  without  any  intermediate  steps,  from  a  plant  with  colored  flowers  to  a  pure  white 
variety;  which  change,  for  want  of  a  better  term,  we  may  call  "spontaneous.  "  Such  may  be 
aptly  termed  "negative"  varieties,  since  their  pecularity  is  due  rather  to  an  absence  of  their 
normal  color  than  to  the  presence  of  white. 

It  is  nothing  uncommon  to  see,  in  many  species,  a  gradual  change  from  a  brightly  colored 
individual,  through  successive  lighter  and  lighter  ones,  until  a  pure  white  is  reached.  This 
may  be  very  well  seen  in  Hepatica  triloba,  which  comprises  individuals  of  every  shade  from 
dark  purple  to  white;  or  in  Polygala  sanguinea,  in  which  we  find  the  same  gradual  change 
from  a  dark  red.     *     *     * 

The  following  list  of  "albinos"  is  made  from  specimens  collected  during  the  last  three 
years.  Vemonia  novaboracensis  W'illd.,  Lappa  officinalis  var.  major,  Lobelia  syphiliiica 
Linn.,  Epiphegus  virginiana,  Verbena  hastata  Linn  ,  Asclepias  incarnata  Linn.,  TrifoHum 
pratense  Linn.,  and  Brunella  vulgaris  Linn.  Both  Gentiana  crinifa  Froel.  and  Lobelia 
cardinalis  Linn,  have  been  reported  to  nie  as  having  produced  at  times  albino  forms,  but  I 
have  never  seen  them  personally.     *     *     * 

Now,  in  the  first  place,  not  only  does  the  flower  show  the  characteristic  absence  of  color 
but  the  leaves,  stem,  and,  in  fact,  the  entire  plant,  are  invariably  of  a  lighter  green;  and  if 
any  red  or  green  should  be  normal  to  the  stem  (which  is  often  the  case)  this  will  also  be  of  a 
lighter  shade.     *     *     * 

Secondly,  if  we  have  under  consideration  a  plant  which  commonly  is  known  to  have 
juice  of  an  acid  or  peculiar  taste,  this  is  generally  more  or  less  absent  in  the  albino  form,  and 
sometimes  is  eliminated  entirely.  Darwin  has  noted  this  fact,  and,  in  commenting  upon  it, 
says  that  honey  bees  evidently  are  aware  of  it,  for  they  perforate  the  calyx  and  corolla  of 
the  white  Aconitum  napcllus,  to  get  at  the  nectaries,  but  will  not  so  do  with  the  colored 
ones.     *     *     * 

It  has  often  been  urged  that  these  albinos  are  mere  "sports"  of  Nature,  with  nothing 
constant  about  them;  their  peculiarities  due,  often,  to  growing  in  the  shade,  etc.  In  fact, 
that  it  is  a  condition  due  to  bleaching  or  insufficient  sunlight,  and  that  there  is  nothing 


76     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 

inherent  in  the  constitution  of  the  plant.  Fortunately  I  have  been  able  to  test  this.  In 
the  case  of  Lobelia  syphilitica  I  first  found  the  plants  in  the  shade  of  some  rather  thick 
underbrush,  in  the  month  of  September.  This  growth  was  cut  down  the  following  spring, 
and  the  place  opened  to  the  full  glare  of  the  sun.  This  was  done  three  years  ago,  yet,  every 
autumn  since,  the  plants  have  either  reappeared,  or  else  left  offspring  which  have  inherited 
their  albino  nature.  This  shows  them  not  only  constant  in  their  peculiarities,  but  also  that 
these  are  bred  in  the  plant  and  capable  of  inheritance.  Epiphcgus  virginiana  and  Brunella 
vulgaris  offer  the  same  proofs.  Nor  has  the  influence  of  locality  much  or  anything  to  do 
with  it,  for  a  plant  of  the  Lobelia  syphilitica  (with  white  flowers),  which  originally  grew  in  a 
swamp,  was  transplanted  to  a  dry  garden  a  mile  or  more  away,  yet  came  up  and  blossomed 
white  the  next  year. 

Let  us  now  see  what  the  experience  gained  in  cultivation  of  white  varieties  can  tell  us. 
Perhaps  the  Japanese  have  brought  the  art  of  eliminating  color  from  plants  to  the  greatest 
perfection.  Scores  of  species  and  genera  have  been  by  them  variegated  in  the  most  peculiar 
manner.  But  this  is  never  constant  in  this  country,  but  after  a  while  always  tends  to 
revert  to  the  primitive  color  again.     *     *     * 

As  there  seems  to  be  no  tendency  to  reversion  in  these  natural  albinos,  they  might  per- 
haps be  made  permanent  varieties  and  be  valuable  on  that  account.  No  doubt  this  per- 
manence is  due  to  the  change  being  sudden,  leaving  no  trace  of  color  by  intermediate  steps, 
while  in  cultivated  examples  the  white  has  generally  been  obtained  by  a  gradual  selection  of 
less  and  less  darkly  colored  ones,  and  hence  there  would  be  a  greater  tendency  to  reversion 
back  through  these  steps  again. 

From  the  foregoing  it  is  clear  that  retrogressive  white-flowered  derivatives 
were  seen  to  arise  from  colored  parental  forms  by  saltations  in  which  the  color 
of  the  flower  behaved  as  a  unit-character,  and  furthermore  that  the  new  color- 
less forms  were  constant  and  were  not  affected  by  environmental  conditions. 
These  ideas  in  fact  underlie  an  important  part  of  the  mutation  theory,  and  the 
statements  by  Dr.  Hollick  constitute  the  unexpanded  thesis  of  the  saltatory 
behavior  of  one  set  of  characters.  Not  the  least  interesting  part  of  the  dis- 
cussion is  that  which  in  effect  recognizes  the  fact  that  many  white-flowered 
and  white-leaved  forms  in  cultivation  represent  the  extremes  of  fluctuating 
variations,  and  that  when  artificial  selection  lapses  these  forms  gradually 
return  to  a  normal  mean ;  also  that  some  species,  such  as  Hcpatica  triloba,  may 
also  have  an  extremely  wide  range  of  fltictuating  variability  as  to  the  flower- 
colors. 

A  dim  realization  of  the  importance  of  sports  and  of  the  necessity  for  some 
other  hypothesis  besides  natural  selection  to  account  for  the  existence  of  all 
living  plants  seemed  to  be  widespread  among  botanists  about  the  time  De  Vries 
began  the  investigations  which  have  led  to  his  conclusions  as  to  the  influence 
of  mutations.  An  interesting  forecast  as  to  the  matter  was  made  by  the  late 
Thomas  Meehan  (Meehan,  1882).     He  said: 

The  conclusion  I  have  been  forced  to  is  that  the  odd  forms  we  often  find  in  nature  are  not 
necessarily  hybrids,  but  are  as  likely,  if  not  more  likely,  to  be  the  outgrowth  of  some  internal 
law  of  form  with  which  we  are  as  yet  unacquainted.  That  they  do  not  often  perpetuate 
themselves  is  \not^  remarkable  when  we  remember  that  of  thousands  of  seeds  produced  on 
anyone  tree  but  a  small  percentage  ever  gets  a  chance  to  form,  and  of  those  which  do  sprout, 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OF   THE    OENOTHERAS. 


/  / 


again  but  a  small  percentage  survives  to  become  bearing  trees.  As  the  number  of  trees 
reproducing  the  general  features  of  the  original  may  be  a  hundred  to  one  of  the  more  strik- 
inglyaberrant  forms,  we  may  see  that  though  individual  instances  may  be  common,  we  are 
never  likely  to  meet  many  trees  of  one  stamp.  Once  in  a  while  an  individual  tree  may  find 
itself  in  a  situation  favorable  to  the  preservation  of  a  number  of  seedlings,  which  might 
endure  until  again  reproductive;  in  such  cases  a  marked  variety  may  originate  and  make 
its  way  over  the  earth. 

I  have  often  thought  it  probable  that  in  time  a  few  individuals  of  these  suddenly  intro- 
duced forms  might  again  leap  into  new  features,  and  then  if  they  should  be  able  to  sustain 
themselves  we  should  have  new  species  quite  independently  of  any  principle  of  natural 
selection. 

The  negative  implied  by  the  context  is  supphed  in  brackets  and  indicated 
by  italics. 

The  above  are  by  no  means  to  be  the  earliest  formulations  of  the  hypothet- 
ical conclusion  as  to  the  importance  of  sports  in  phylogeny.  Among  others 
which  might  be  mentioned  the  statement  by  Kerner  in  1869  is  worthy  of 
citation.     He  says: 

Die  Arten  die  uns  gegeniibertreten,  sind  nur  Stadien  and  haben  als  solclie  zwar  fiir  eine 
gewisse  Zeit  Konstanz,  konnen  sich  aber  friiher  oder  spater  in  andersgeformte  Arten  auflo- 
sen.  Thatsache  ist  es,  dass  alle  Pflanzen  friiher  oder  spater  einmal  vereinzelte  Abarten, 
d.  i.  S])r6sslinge  zu  erzeugen  in  Stande  sind,  welche  in  ihren  Merkmalen  von  der  Mutterart 
abweichen,  und  zahlreichen  Erscheinungen  drangen  uns  zu  der  Annahme,  dass  unter  dem 
zusammentreffen  giinstiger  Bedingungen  solche  Abarten  die  Ausgangspunkte  neuer  Arten 
werden. 

Among  other  recent  observations  is  that  of  Schaflfner  (1906)  concerning 
a  derivative  of  Verbena  stricta  which  bore  pinkish-white  instead  of  the  deep 
purple  flowers  of  the  species.  The  aberrant  form  was  present  in  many  thou- 
sands of  individuals  among  the  typical  specimens,  with  which  it  seemed  to  be 
in  successful  competition.  Tests  have  not  yet  been  made  to  determine 
whether  or  not  it  does  not  readily  cross  with  the  parental  form,  or  whether  it 
is  dominant  or  recessive  with  regard  to  the  more  important  differentiating 
character.  In  any  case,  however,  its  mutative  origin  is  based  upon  evidence 
similar  to  that  which  zoologists  seem  willing  to  accept  as  proof  of  mutation  in 
animals,  and  its  successful  maintenance  against  parental  competition  is  an 
established  fact. 

Many  of  the  observations  brought  forward  by  various  authors  concern 
species  which  have  been  long  under  cultivation,  and  so  many  facts  reported 
from  horticultural  material  have  been  found  to  be  unguarded  against  errors 
affecting  purity  of  lineage  that  much  prejudice  exists  with  respect  to  data 
obtained  from  domesticated  forms.  So  far  as  the  effects  of  actual  tillage  or 
domestication  are  concerned,  these  prejudices  are  without  foundation,  since 
at  the  present  time  no  well-founded  evidence  exists  to  show  that  cultural  con- 
ditions have  ever  produced  alterations  which  were  strictly  and  continuously 
inheritable  under  environic  conditions  other  than  those  by  which  the  altera- 


7$     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OP  THE   OENOTHERAS. 

tions  were  induced,  although  vague  statements  and  erroneous  generahzations 
to  the  contrary  are  current.  The  possibiHty  of  such  results  is  to  be  by  no 
means  denied,  but  if  this  view  is  to  be  upheld  it  must  be  supported  by  some 
evidence  based  upon  actual  experimentation.  Vicinism,  the  somatic  multipli- 
cation of  bud-sports  and  extreme  variants,  and  the  confusion  of  closely  related 
elementary  species  form  the  basis  for  the  greater  number  of  positive  assertions 
as  to  the  effects  of  cultivation,  and  it  is  necessary  to  examine  all  facts  bearing 
upon  the  lineage  of  supposedly  new  forms  with  the  greatest  care  before  their 
aspect  or  behavior  may  be  taken  as  evidence  upon  phylogenetic  problems. 

An  extended  consideration  of  the  phenomena  of  mutability  has  led  the 
authors  of  this  paper  to  entertain  grave  doubts  as  to  the  practical  value  of 
the  conception  of  periodicity.  De  Vries  throughout  his  writings  has  favored 
the  assumption,  on  purely  theoretical  grounds,  that  a  species  must,  at  certain 
times  in  its  history,  be  in  a  mutable  condition,  while  in  other  periods  of  great 
length  all  of  the  progeny  come  true  to  the  type,  subject  only  to  fluctuating 
variability. 

It  is  quite  possible  that  any  given  form  may  give  off  derivatives  steadily 
for  centuries,  but  none  of  these  being  fitted  to  survive,  no  trace  of  this  activity 
remains.  A  change  in  the  character  of  the  mutants  originated,  or  newly 
found  conditions  met  with  in  comparatively  short  migrations,  might  readily 
give  an  appearance  of  the  beginning  of  a  mutative  period,  unless  the  matter 
had  been  tested  by  pedigree-cultures. 

With  regard  to  the  actual  succession  of  generations  two  aspects  of  the  case 
present  themselves.  In  one  case  it  may  be  supposed  that  an  annual  species 
might  have  a  limited  distribution,  and,  occupying  small  areas,  might  be  repre- 
sented by  only  a  few  individuals  every  year.  Combined  with  these  circum- 
stances, the  proportion  of  mutants  produced  by  the  species  might  be  so  small 
that  it  would  be  possible  to  examine  every  specimen  for  a  hundred  years 
without  meeting  a  single  atypic  individual.  The  conclusion  would  then  be 
reached  that  this  was  included  in  the  immutable  period.  Then  if  the  number 
of  individuals  grown  and  examined  in  the  next  ten  years  numbered  as  many 
millions,  and  some  mutants  were  found,  the  line  of  reasoning  followed  would 
lead  to  the  fallacious  assumption  that  a  mutable  period  had  begun,  or  rather 
that  some  physiological  change  had  ensued  in  consequence  of  which  mutations 
were  occurring.  These  might  as  readily  be  seen  during  the  first  year  of 
observation,  however,  if  enough  individuals  had  been  grown. 

It  appears,  therefore,  that  the  real  state  of  affairs  is  better  represented  by 
the  phrase  "frequency  of  mutation,"  by  which  is  expressed  the  number  of 
individuals  which  must  be  grown  to  furnish  one  mutant,  and  which  is  nearly 
identical  with  "the  coefficient"  of  mutability.  If  the  parental  type  pro- 
duces more  than  one  mutant,  the  frequency  of  these  may  vary  widely  from 
each  other. 


MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OF  THE   OENOTHERAS.      79 

Thus  of  the  great  number  of  mutant  individuals  originated  by  0.  lamarck- 
iana  but  few  have  been  ruhrinervis,  still  a  smaller  number  represented  gigas, 
while  the  combined  observations  at  New  York  and  Amsterdam  have  failed  to 
bring  hrevistylis  within  the  range  of  the  cultures,  this  form  having  arisen  at 
Hilversum,  where  it  is  still  in  existence.  To  secure  mutants,  or  to  allow  a 
species  to  display  its  possible  derivatives,  the  long  stretch  of  centuries  may  be 
shortened  and  as  many  individuals  examined  in  a  seedling  stage  in  a  single 
season  as  might  naturally  mature  in  twenty  decades.  The  real  point  to  be 
attained  is  to  secure  sufhcient  individuals  to  include  the  proportionate  num- 
ber of  all  the  derivatives. 

ADDITIONAL    EVIDENCE    AS   TO    THE    DISTRIBUTION    AND   OCCURRENCE 

OF   LAMARCK'S  EVENING-PRIMROSE. 

The  records  and  material  cited  and  examined  show  that  a  large-flowered 
evening-primrose,  which  can  be  identified  with  nothing  so  closely  as  with 
0.  lamarckiana,  had  found  its  way  into  the  gardens  near  Haarlem,  Holland,  as 
early  as  1756,  several  years  before  0.  grandiflora  was  brought  from  its  habitat 
in  Alabama  to  England  in  1778.  Also  that  an  evening-primrose  with  large 
flowers  was  seen  on  the  sand  banks  north  of  Liverpool,  England,  in  1806,  on 
the  coast  of  Somerset  in  1837,  and  has  been  cited  and  described  by  various 
authors  in  1845,  1851,  1855,  i860,  and  continuously  since  1892.  Plants 
grown  from  seeds  obtained  from  the  region  in  question  proved  to  be  0.  lamarck- 
iana with  two  of  its  mutants,  rubrincrvis  and  lata.  Not  only  did  this  con- 
firm the  supposition  as  to  the  long  history  of  lamarckiana,  but  it  also  showed 
that  rubrincrvis  had  successfully  maintained  itself  in  competition  with  the 
parental  form,  while  lata  had  become  capable  of  self-fertilization,  the  only 
instance  on  record. 

THE  MUTANTS  AND  MUTABILITY  OF  LAMARCK'S  EVENING-PRIMROSE. 

The  observations  begun  by  De  Vries  upon  0.  lamarckiana  in  1884  and 
continued  upon  purely  pedigreed  material  by  himself  until  the  present  time, 
and  the  cultural  experiments  described  in  this  paper,  demonstrated  a  very 
high  frequency  of  mutation  in  this  species  and  also  that  the  mutative  depart- 
ures show  a  great  diversity  when  compared  with  other  forms.  In  addition  to 
the  derivatives  which  constitute  the  greater  proportion  of  the  atypic  offspring 
gigas  has  recently  reappeared  in  an  example  in  New  York,  being  one  of  the 
rarer  forms.  Then  the  sudden  appearance  of  a  totally  new  type  among  the 
mutants  leads  at  once  to  the  confirmation  of  the  suggestion  that  certain 
mutants  may  appear  so  infrequently  that  they  might  not  be  expected  more 
than  once  in  a  million  individuals  and  consequently  would  not  be  seen  except 
at  long  intervals. 

Of  the  known  mutants  alhida,  oblonga,  gigas,  nanella,  lata,  and  scintillans 
were  found  in  the  pedigreed  cultures  described  in  this  paper,  and  with  greatly 


8o     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS    OP   THEl    OE;NOTHE;rAS. 

varying  frequency.  In  one  culture  the  coefficient  of  mutability  of  oblonga 
was  pushed  slightly  beyond  the  limit  hitherto  assigned  to  it  by  De  Vries  and 
it  was  found  to  constitute  54  per  cent  of  the  atypic  derivatives.  A  survey  of 
all  of  the  results  obtained  in  the  New  York  Botanical  Garden  since  1902  does 
not  justify  the  assertion,  however,  that  any  change  in  the  frequency  of  mutation 
has  been  induced  by  cultural  conditions,  and  the  only  really  new  feature 
noticed  consisted  in  the  appearance  of  a  new  mutant,  which,  however,  was  so 
unsuited  to  the  climatic  conditions  that  it  perished  before  reaching  maturity, 
an  example  of  the  eliminative  action  of  selection  in  the  determination  of  what 
mutants  shall  survive  and  what  shall  perish. 

Cultures  made  from  seeds  obtained  from  merchants  did  not  show  a  fre- 
quency of  the  mutants  above  1.5  per  cent  in  any  instance,  which  is  much  less 
than  that  observed  in  the  strain  growing  at  Hilversum.  The  sources  from 
which  the  seeds  were  obtained  indicate  that  the  species  is  in  a  state  of  muta- 
bility in  various  parts  of  the  world,  and  that  it  has  probably  been  so  for  some 
time.  The  low  frequency  of  mutation  may  have  been  due  in  part  to  unsuit- 
able methods  of  cultivation,  in  consequence  of  which  the  parental  individuals 
were  badly  nourished. 

0.  hrevistylis  was  brought  to  maturity  and  found  to  agree  in  all  characters 
with  specimens  grown  in  Amsterdam.  This  form  differs  from  the  parental 
type  chiefly  by  the  retrogressive  character  of  a  shortened  style,  and  by  gen- 
eral observation  this  organ  shows  a  wider  range  of  fluctuation  about  its  average 
than  the  same  organ  in  the  parent,  although  no  exact  examination  was  made. 
Despite  the  fact  that  this  type  produces  comparatively  few  seeds  it  was  found 
by  De  Vries  among  the  individuals  of  the  parental  type  in  the  original  locality 
in  which  mutation  in  lamarckiana  was  observed  in  1886,  and  it  has  since  held 
its  own  in  competition  with  the  parental  type,  in  a  manner  demonstrative  of 
the  fact  that  it  is  possible  for  a  mutant  to  survive  in  competition  with  existing 
forms. 

O.  lata  grown  from  seeds  obtained  from  Professor  De  Vries  and  from  indi- 
viduals arising  in  the  pedigreed  strains  in  New  York  was  seen  to  remain  con- 
stant to  its  characters.  A  few  tests  had  been  made  both  in  New  York  and 
Amsterdam,  but  no  fertilization  was  secured  with  its  own  pollen,  although  some 
of  it  appeared  structurally  perfect  in  a  microscopic  examination.  0.  lata 
appearing  in  a  culture  from  seeds  of  0.  lamarckiana  from  England,  however, 
included  some  individuals  which  likewise  produced  pollen,  and  also  showed 
the  hitherto  unknown  capacity  of  being  fertilized  by  it.  The  seeds  obtained 
in  this  way  gave  rise  to  a  progeny  which  showed  only  the  constituents  usually 
found  in  a  progeny  of  this  plant  when  fertilized  by  lamarckiana. 

O.  scintillans  grown  directly  from  its  own  seeds,  and  also  as  it  appeared  as  a 
new  mutant  in  the  cultures,  was  constant  in  its  ever-sporting  character.  Seeds 
of  this  species  purely  fertilized  gave  rise,  in  addition  to  the  type,  to  a  few 


MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE)    OENOTHERAS.      8 1 

individuals  of  oblonga  and  lamarckiana,  in  addition  to  an  unrecognizable  form, 
the  representative  of  which  did  not  reach  maturity.  Purely  fertilized  seeds  of 
this  form  have  also  been  seen  to  give  rise  to  lata  and  nanclla. 

O.  oblonga  was  grown  directly  from  its  own  seeds  and  also  appeared  in  all 
of  the  cultures  of  the  parental  type  which  were  spread  sufficiently  to  include 
its  coefficient  of  mutability.  It  appears  to  be  constant  in  its  characters  and 
is  strikingly  differentiated  at  a  very  early  stage  of  its  development. 

O.  albida  was  also  grown  directly  from  its  own  seed  and  appeared  in  the 
pedigreed  cultures  of  lamarckiana.  It  appeared  to  coincide  in  all  of  its 
characters  with  the  descriptions  given  by  De  Vries. 

The  single  test  which  was  made  of  the  effect  of  crossing  the  parental  type, 
lamarckiana,  with  one  of  its  mutants,  rubrinervis,  gave  results  in  accordance 
with  the  conclusions  of  De  Vries  in  this  matter.  Rubrinervis  is  a  mutant  of 
supposedly  low  frequency  of  occurrence,  since  it  has  not  been  found  in  any 
pedigreed  culture  for  several  years.  It  may  be  regarded  as  a  progressive 
derivative  of  the  parental  type  and  presumably  carries  all  of  the  lamarckiaim 
qualities  in  a  latent  condition.  On  the  other  hand,  the  strain  of  lamarckiana 
with  which  it  was  crossed,  and  from  which  it  was  derived  several  generations 
back,  must  bear  the  characters  which  gave  rise  to  this  form  as  a  mutant.  The 
two  plants  must  therefore  carry  the  same  characters,  but  some  are  latent  in 
one  and  some  in  the  other. 

The  origin  of  species  by  hybridization,  which  has  been  a  well -recognized 
phenomenon  for  half  a  century,  receives  further  exemplifications  in  crosses 
between  0.  cruciata  and  0.  lamarckiana.  The  fixed  forms  which  may  be  pro- 
duced by  this  cross  have  been  cultivated  in  European  gardens  for  some  time 
under  the  names  of  0.  cruciata,  0.  cruciata  varia,  and  0.  biennis  cruciata.  The 
list  of  species  which  have  originated  by  hybridization  is  a  long  one  and  need 
not  be  recounted  here.  Singularly  enough,  this  fact  seems  to  be  unknown  to 
workers  who  carry  on  extensive  operations  in  plant  breeding.  In  the  instances 
cited  in  this  paragraph,  the  most  remarkable  feature  is  that  a  fixed  form  or 
new  species  may  be  produced  by  the  hybridization  of  two  parental  types  of 
comparatively  high  frequency  of  mutability.  It  seems  also  pertinent  to  again 
sound  a  warning  against  the  continued  practice  of  taxonomists  of  ascribing 
hybrid  origin  to  species,  with  characters  apparently  intermediate  between  two 
neighboring  and  known  forms.  Numerous  hybrids  of  known  origin  in  the 
New  York  Botanical  Garden  have  been  submitted  to  competent  taxonomists, 
and  in  no  single  instance  has  a  correct  diagnosis  of  the  parentage  been  made. 

The  fertilization  of  the  eggs  of  lamarckiana  from  the  pollen  of  rubrinervis 
resulted  in  a  single  observation  in  a  progeny  which  contained  71  per  cent  of 
the  latter,  or  mutant,  type.  This  is  not  a  universal  result,  however,  since  the 
proportion  has  been  found  to  vary  from  19  to  74  per  cent.  The  pollination  of 
0.  biennis  hy  the  mutant  described  in  this  paper  gave  a  progeny  in  the  first 


82     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OF   THE)   OENOTHERAS. 

generation  which  conformed  to  the  biennis  type.  The  same  result  is  true  of 
the  reciprocal  cross.  It  remains  to  be  seen  whether  the  mutant  characters  are 
merely  recessive  in  this  combination,  or  whether  a  condition  of  latency  has 
been  assumed  in  which  the  mutant  appears  only  in  its  customary  frequency. 

The  evening-primroses,  then,  afford  the  following  exact  evidence  as  to  the 
effects  of  intercrossing :  Many  of  the  mutants  when  crossed  with  the  parental 
form  give  progeny  composed  of  the  parental  form  and  the  mutant  in  the  first 
generation.  In  the  case  of  gigas  and  rubrinervis  the  proportion  of  the  mutant 
in  the  progeny  is  generally  much  higher  than  the  parental  type.  Rubrinervis 
when  grown  in  close  contiguity  to  the  parental  form  showed  only  a  small  pro- 
portion of  crosses,  and  the  newly  discovered  mutant  of  0.  biennis  also  gave 
but  a  small  proportion  of  crosses  when  exposed  to  danger  of  hybridization 
with  the  parent.  Lastly  it  is  to  be  pointed  out  that  0.  brevistylis,  which  pro- 
duces but  few  seeds,  and  which  is  recessive  when  crossed  withO.  lamarckiana, 
has  been  in  continuous  existence  for  20  years,  perhaps  much  longer,  at  Hil- 
versum,  where  it  is  in  direct  competition  and  danger  of  being  hybridized  with 
the  parental  form.  Experiences  in  the  guarded  cultures  seem  therefore  to  be 
confirmed  by  an  analysis  of  natural  conditions.  Swamping  of  new  forms,  by 
intercrossing,  is  a  specter  which  looms  vast  and  shadowy  across  the  visions  of 
writers  of  a  speculative  habit,  but  at  the  present  moment  it  is  believed  that  no 
evidence  carefully  tested  by  modem  methods  is  available  to  justify  the  hypoth- 
esis, so  far  as  plants  are  concerned. 

The  demonstration  of  the  greater  variability  of  the  mutants  as  compared 
with  the  parent  form  seems  to  be  complete,  and  this  relation  is  one  of  consid- 
erable practical  and  theoretical  importance.  The  greater  variability  of  phylo- 
genetically  new  characters  as  compared  with  older  ones  is  not  wholly  new  and 
unique,  though  we  believe  our  studies  in  1904  gave  the  first  statistical  support 
to  the  thesis  when  stated  in  this  form.  Dar\Ndn's  (1859,  Ch.  II)  view  that 
characters  which  distinguish  species  are  more  variable  than  those  that  sepa- 
rate genera,  and  that  varietal  are  more  variable  than  specific  characteristics, 
gives  the  first  recognition  of  this  principle,  if  we  take  into  account  that  accord- 
ing to  his  conception  of  the  relation  of  these  several  form-groups  the  variety  is 
an  incipient  species  and  the  species  an  incipient  genus.  Field  (1898)  worked 
out  the  relation  between  a  varietal  and  a  specific  characteristic  in  the  wings 
of  a  lepidopterous  insect,  giving  a  confirmation  of  Darwin's  rule,  but  he  did 
not  relate  his  facts  in  any  way  to  the  relative  age  of  the  characters  studied. 
It  is  probable  of  course  that  the  varietal  characteristic  was  a  more  recent 
acquisition  than  the  specific,  and  this  would  make  his  results  accord  well  with 
the  observations  of  the  authors  on  Oenothera. 

To  the  student  of  evolution  who  wishes  to  institute  pedigree-cultures,  the 
relation  here  demonstrated  should  prove  of  much  practical  value,  for  it  matters 
not  whether  he  desires  to  study  the  possibility  of  fixing  extreme  fluctuations, 


MUTATIONS,    VARIATIONS,    AND   REI^ATIONSHIPS   OF   THE   OENOTHERAS.      83 

or  whether  he  is  engaged  in  tracing  the  character,  frequency,  and  causes  of 
mutation  a  preHminary  statistical  investigation  will  aid  in  the  choice  of  mate- 
rial best  suited  to  his  purpose.  It  is  obvious  that  the  great  variabilitv  of  any 
elementary  species  would  disfavor  it  as  research  material  for  the  study  of 
mutations  rather  than  indicate  its  desirability,  as  the  wide  range  of  its  fluctua- 
tions would  suggest  the  likelihood  of  its  being  a  recent  mutant  rather  than  a 
mutating  species.  A  more  promising  procedure  with  respect  to  any  variable 
form  would  seem  to  be  to  use  the  most  nearly  related  or  contiguous  species 
having  a  low  degree  of  variability  in  the  hope  of  thus  hitting  upon  the  parental 
type  from  which  the  variable  form  originated.  The  student  of  pedigree- 
cultures  frequently  receives  letters  from  the  kindly  disposed,  calling  his  atten- 
tion to  strikingly  variable  forms  which  are,  on  the  ground  of  their  variability, 
conceived  to  be  especially  valuable.  It  is  to  be  hoped  that  in  the  future  more 
attention  will  be  given  to  constant  forms  having  one  or  more  variable  near 
relatives. 

Whether  the  relative  variability  of  old  and  new  structures  could  likewise  be 
used  by  the  morphologist  as  a  new  criterion  for  working  out  lines  of  descent  it 
would  be  somewhat  reckless  to  say  at  present,  but  an  important  field  for 
research  is  indicated.  Are  forms  which  are  obviously  more  recent  at  the  same 
time  more  variable  than  the  older?  Are  the  structures  characteristic  of  the 
Orchidaceae,  for  instance,  more  variable  than  those  of  more  generalized  mono- 
cotyledons, and  those  of  the  Compositae  more  variable  than  those  of  other 
dicotyledons?  Are  monocots  on  the  whole  less  variable  than  the  dicots? 
While  not  wishing  to  press  the  application  of  the  principle  too  far,  it  may  be 
pointed  out  that  one  feature  of  the  investigation  here  reported  seems  to  bear 
directly  upon  the  phylogenetic  significance  of  the  greater  variability  of  new 
characters.  The  characteristic  feature  of  the  Oenothera  bud  is  the  hypan- 
thium.  It  doubtless  represents  one  of  the  most  fundamental  mutations  in  the 
history  of  the  genus.  It  accords  well  with  the  hypothesis  here  offered  to  find 
that  the  length  of  the  hypanthium  is  in  every  species  the  most  variable  part 
of  the  bud.  The  great  excess  in  its  variability  over  that  of  other  parts  of  the 
buds  is  made  strikingly  apparent  by  comparing  the  average  variability  of  the 
several  parts  of  the  buds  in  all  the  species  studied.  The  average  coefficients  of 
variability  are  as  follows,  in  order  of  their  magnitude :  Length  of  hypanthium, 
14.97;  length  of  cone,  9.56;  length  of  ovary,  8.66;  thickness  of  cone,  7.85; 
thickness  of  ovary,  6.30 ;  thickness  of  the  hypanthium,  5.83.  The  consistency 
of  these  results  make  it  certain  that  the  great  variability  of  the  hypanthium 
originated  with  the  origin  of  the  hypanthium. 

The  fact  that  a  new  species  is  more  variable  than  an  old  one  should  also 
give  to  the  phytogeographer  a  method  of  investigating  the  exact  status  of  an 
endemic  species.  If  it  presents  a  case  of  initial  endemism  it  should  be  more 
variable  than  the  nearly  related,  wide-ranging,  and  hence  presumably  older 


84     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OI^   THE   OENOTHERAS. 

species.  If  on  the  other  hand  it  is  a  local  relict  of  an  old  species,  which  once 
had  a  wide  distribution,  it  should  present  less  variability  than  its  wide  ranging, 
presumably  newer  relative.  The  only  satisfactory  evidence  heretofore  avail- 
able for  determining  a  point  of  this  kind  has  been  drawn  from  paleontological 
sources. 

To  the  metaphysician  who  is  striving  to  picture  to  himself  the  laws  of  vital 
motion,  the  results  of  these  statistical  studies  will  give  an  altogether  new  view 
of  the  relation  between  fluctuation  and  mutation.  Rosa  (1899)  has  published 
a  treatise  in  which  he  contends  that  there  is  a  progressive  lessening  of  varia- 
bility during  the  progress  of  evolution,  and  he  sees  as  the  certain  result  of  this 
process  the  ultimate  extinction  of  life.  Plate  (1904)  takes  issue  with  Rosa  on 
this  point  and  maintains  that  there  is  no  such  law.  The  demonstration  that 
there  is  greater  variability  in  new  species,  coupled  presumably  with  a  decrease 
in  variability  as  the  species  grows  older,  indicates  that  both  Rosa  and  Plate 
are  partly  right.  It  is  now  seen  that  although  the  individual  species  may 
decrease  in  variability  as  it  grows  older,  this  decrease  is  compensated  for  every 
time  a  new  species  springs  into  existence.  Instead  of  mutations  being  the 
cumulative  results  of  ever-increasing  fluctuation,  they  appear  now  as  an  initial 
process  of  which  fluctuation  is  in  part  an  after  effect. 

A  detailed  description  of  but  a  few  of  the  species  of  Oenothera  have  been 
given.  Seeds  and  material  have  been  brought  in  from  all  parts  of  America 
east  of  the  Rocky  Mountains,  and  a  consideration  of  this  material  gives 
foundation  for  the  conclusion  that  under  the  name  biennis  is  included  a  swarm 
of  elementary  species  very  closely  related,  but  easily  distinguishable  when 
grown  side  by  side,  having  no  intergrading  forms. 

Even  the  imperfect  knowledge  we  possess  of  the  species,  owing  to  the 
scarcity  of  collections  made  of  them,  shows  that  they  overlap  in  distribution 
in  the  most  haphazard  manner.  In  the  same  way,  the  elementary  species 
comprised  within  cruciata  interlock.  In  discussing  the  principles  that  govern 
distribution,  it  is  to  be  admitted  that  in  accordance  with  their  usual  practice 
many  systematists  would  not  allow  specific  rank  to  the  forms  in  question,  so 
closely  are  they  related.  It  will  be  in  order,  therefore,  to  take  up  the  next 
grouping  of  interest,  that  of  biennis,  and  the  allied  species  of  oakesiana,  parvi- 
flora,  7nuricata,  and  cruciata,  which  were  accepted  as  varieties  of  biennis  30 
years  ago,  but  which  are  accorded  specific  rank  at  the  present  time.  It  is 
found  that  biennis  blankets  nearly  the  entire  range  of  these  forms,  and  also 
that  the  ranges  of  these  forms  overlap  among  themselves.  It  is  to  be  seen, 
therefore,  that  no  matter  what  range  of  opinion  may  enter  into  the  discussion 
as  to  the  rank  of  the  forms  of  the  evening-primroses  of  the  eastern  part  of 
North  America,  the  thesis  that  the  most  nearly  related  species  do  not  overlap 
in  distribution  is  unsupported. 


MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OF   THE   OENOTHERAS.     85 

A  condition  similar  to  the  above  has  been  found  in  Crataegus  by  Sargent 
&  Peck,  who  say  (1906) : 

The  peculiar  tendency  of  Crataegus  to  flock  together  is  strikingly  illustrated  in  our  terri- 
tory. It  is  rare  to  find  any  large  area  occupied  by  a  single  species.  Where  many  thorn 
trees  and  bushes  grow  together  there  are  usually  many  species.  A  remarkable  example  of 
this  kind  is  found  in  a  narrow  strip  of  pasture  land  bordering  the  Erie  Canal  near  Menands. 
Here  10  species  are  growing  in  an  area  of  about  i  acre.  The  closest  condensation  of  num- 
erous species  that  I  have  seen  anywhere  is  near  Albia,  where  9  of  our  native  species  are 
growing  in  a  kind  of  irregular  row  along  the  west  bank  of  the  Wyantskill  Creek.  The 
length  of  the  row  is  about  100  feet.  It  is  worthy  of  remark  that  3  of  these  species,  Cratae- 
gus fereiitaria,  C.  rhombifolia,  and  C.  succulenta,  belong  to  the  group  Tomentosae.  Such 
close  associations  of  niembers  of  a  single  group  as  this  one  at  Lansingburg  are  very  signif- 
icant and  when  more  fully  understood  may  possibly  throw  some  light  on  the  interesting 
problem  of  the  development  of  species. 

Other  students  of  the  thorns  have  found  similar  conditions  with  strains 
which  were  found  to  be  distinct  and  hereditary  in  pedigreed  cultures.  Here, 
again,  if  a  conservative  view  is  upheld  and  specific  rank  is  not  accorded  these 
clearly  separable  elementary  species,  a  similar  association  of  the  closely 
related  group  species  may  be  found.  A  similar  condition  among  the  opuntias 
has  been  noted  by  the  author.  These  facts  make  it  conclusive  that  the 
assumption  that  the  nearly  related  or  most  nearly  related  species  of  a  group 
do  not  occupy  the  same  habitat  is  untenable. 

Two  bud-sports  of  different  types  as  to  origin  were  found  during  the 
summer  of  1905  and  tested  as  to  their  hereditary  qualities. 

The  vegetative  mutant  branch  of  ammophila  was  seen  to  give  rise  to  a 
progeny  which  came  true  to  the  characters  of  the  branch  producing  the  seeds 
from  which  they  were  grown,  and  the  reversion,  if  it  is  such,  may  be  consid- 
ered complete.  If,  as  suggested,  ammophila  is  a  hybrid  and  the  sport  is  a  rever- 
sion to  one  of  the  parents,  then  this  case  may  be  taken  to  agree  with  that  noted 
by  De  Vries  (1900,  p.  86),  in  v.'hich  a  hybrid  of  two  forms  of  Veronica  gave  off 
bud-sports,  which  in  purely  fertilized  seeds  came  wholly  true  to  one  of  the 
parental  types.  In  this  case  the  sport  was  a  reversion  to  the  ancestral  qualities 
which  had  become  latent  when  the  hybridization  was  effected. 

The  second  bud-sport  was  one  in  which  the  recessive  characters  appeared  on 
a  branch  in  the  first  generation  of  a  cross,  and  when  this  was  purely  fertihzed 
showed  only  a  strict  inheritance  of  the  recessive  characters. 

The  facts  offered  by  bud-sports  have  much  significance  as  to  the  localization 
of  mutations,  and  as  to  the  nature  of  the  stimuH  which  set  the  mutatory  pro- 
cesses in  action.  Primary  buds  usually  arise  laterally  from  the  tip  of  the 
growing  point  and  within  a  millimeter  or  two  from  its  apex.  The  saltation, 
of  whatever  character  it  may  be,  occurred  in  a  cell  or  cells  in  the  embryonic 
tissue  while  the  growing  tip  was  moving  upward  a  distance  of  a  millimeter  or 
more.     Much  depends  upon  the  place  where  this  occurs.     If  it  is  possible  for  a 


86     MUTATIONS,    VARIATIONS,    AND   RELATIONSHIPS   OF   THE   OENOTHERAS. 

single  cell  near  the  apex  of  the  growing  point  to  undergo  such  changes  during 
the  rapid  division  that  characterizes  this  region  that  it  gives  rise  to  a  mass  of 
cells  from  which  the  bud  arises,  this  would  favor  the  probability  that  the  salta- 
tory changes  are  due  entirely  to  internal  causes  and  might  not  be  affected  at  all 
by  circumstances. 

Examples  of  sports  are  known  in  which  a  section  of  the  shoot  involving 
several  branches  are  of  an  atypic  character.  DeVries  offers  an  illustration  in 
his  description  of  a  green  branch  on  a  colored  oak  in  which  the  greening  not 
only  affected  the  entire  branch,  but  also  a  section  of  the  member  from  which  it 
arose,  showing  that  the  mutated  protoplast  was  one  which  gave  rise  to  more 
than  the  atypic  branch.  In  other  instances  the  mutation  may  involve  only  a 
longitudinal  section  of  a  branch  or  of  a  member  or  limited  growth  such  as 
inflorescence,  and  this  may  be  carried  to  the  extent  that  even  half  of  a  flower 
may  be  of  an  atypic  character,  and  likewise  similarly  complex  fruits  may  be 
produced.  It  is,  of  course,  admissible  that  in  reversionary  sports,  as  most  of 
these  sectional  variations  are,  the  qualities  that  appear  in  the  sport  are  latent 
in  the  entire  plant  and  need  but  a  slight  stimulus  to  awaken  them,  or  some 
force  to  weaken  the  dominancy  of  the  main  stock.  So  far  as  the  saltation  here 
is  concerned  it  is  one  in  which  any  disturbance  of  the  equilibrium  would  in  all 
probability  have  but  one  result. 

In  the  case  of  the  appearance  of  characters  not  inherited  in  a  latent,  reces- 
sive, or  active  condition,  but  which  constitute  a  progression  or  acquisition,  the 
case  is  different,  and  if  more  than  one  protoplast  is  concerned  in  the  alterations 
ensuing  preliminary  to  the  organization  of  a  bud-sport,  then  the  suggestion 
that  the  alterations  were  the  result  of  stimulation  from  factors  external  to  the 
cell  in  which  the  changes  ensued  would  have  great  force. 

Bud-sports  usually  occur  on  the  lower  part  of  the  main  stem,  where  many 
of  them  lie  dormant  in  sleeping  buds,  and  may  be  awakened  by  decapitation 
of  the  shoot.  It  is  thus  to  be  seen  that  they  belong  to  a  comparatively  early 
stage  in  the  life  of  the  sporophyte.  The  mutations  which  give  rise  to  atypical 
seedlings,  on  the  other  hand,  seem  on  theoretical  grounds  to  be  due  to  changes 
ensuing  in  the  very  closing  stages  of  the  sporophyte,  and  previous  to  the  quali- 
tative or  reducing  divisions  which  form  the  egg,  or  in  the  early  stages  of  the 
pollen  mother-cells.  Here  is  encountered  a  feature  not  yet  recognized  in  bud- 
mutations — that  of  a  fairly  constant  frequency. 

No  evidence  has  yet  been  secured  to  show  that  the  pressure  or  direct  action 
of  environmental  factors  upon  the  vegetative  organs  has  produced  any  per- 
manent inheritable  changes  in  elementary  strains  of  plants.  The  record  of  the 
experiments  in  which  solutions  of  various  kinds  were  injected  into  ovaries 
immediately  previous  to  fertilization  seems  to  point  definitely  to  the  con- 
clusion that  the  bearers  of  the  hereditary  characters  may  be  definitely  and 
directly  affected  by  forces  external  to  the  cell,  however.     The  alterations  thus 


MUTATIONS,    VARIATIONS,    AND    RELATIONSHIPS    OF   THE    OENOTHERAS.      87 

induced  consist  in  the  suppression  of  some  qualities  of  the  parental  form  and 
the  substitution  therefor  of  other  characters  with  which  the  suppressed 
features  are  mutually  exclusive.  The  changes  are  expressed  in  anatomical 
alterations  and  modifications  of  attendant  functions. 

Atypic  derivatives  have  been  secured  in  this  manner  from  two  species, 
Raimannia  odorata  and  Oenothera  biennis,  hy  the  use  of  more  than  one  reagent, 
as  described  in  the  preceding  text.  The  induced  mutants  have  been  tested  to 
the  second  and  third  generation,  with  the  result  that  both  are  found  constant 
in  the  sense  that  they  do  not  intergrade  back  to  the  parental  form. 

The  injection  of  the  solutions  into  the  ovaries  results  in  the  total  destruction 
of  some  eggs  by  the  mechanical  action  of  the  needle  and  of  the  solution  itself, 
while  in  a  few  the  stimulative  or  other  action  is  of  such  nature  that  the  changes 
are  induced  which  result  in  the  modification  of  the  hereditary  characters  and 
of  the  adult  sporophytes  which  carry  them,  while  doubtless  in  most  of  the 
cases  no  action  at  all  resulted.  Of  course  in  some  of  the  unsuccessful  tests  all 
of  the  ovules  were  destroyed  and  fell  off  before  maturity.  While  it  seems 
more  natural  to  assume  that  the  injection  results  in  direct  action  of  the  fluids 
on  the  egg-cells,  yet  the  possibility  is  by  no  means  excluded  that  the  saturation 
of  the  tissues  of  the  ovary  with  the  solution  results  in  the  induced  effect  being 
due  directly  to  the  action  of  the  reagent  upon  the  advancing  pollen-tube  and 
its  contents. 

The  wide  variety  of  substances  used  as  reagents  in  securing  the  alterations  in 
question  makes  it  impossible  to  suggest  the  nature  of  the  changes  induced  in 
the  reproductive  elements,  since  the  actual  effect  may  be  osmotic  or  chemical. 
Neither  may  it  be  said  whether  the  changes  consist  in  disturbances  of  the  pro- 
cessions of  enzymatic  action,  or  actually  modify  the  structure  and  composition 
of  the  cytoplasm  or  nuclear  constituents. 

Mutants  have  thus  been  produced  by  conditions  brought  about  in  an  exper- 
imental way,  and  it  can  not  be  denied  that  similar  action  of  external  forces 
might  occur  unaided  by  man.  The  radioactivity  of  spring  and  rain  water,  or 
the  effects  of  corrosive  or  other  gases  which  are  being  set  free  in  numberless 
places  on  the  earth's  surface,  might  produce  effects  similar  to  these  obtained 
in  the  pedigreed  cultures.  Undue  or  abnormal  secretions  from  the  walls  of  the 
ovary,  from  neighboring  tissues,  or  from'  intruding  foreign  pollen  incapable 
of  accomplishing  fertilization  might  exert  a  similar  influence.  The  stings  and 
lacerations  of  insects  and  animals  and  the  action  of  parasitic  fungi  are  known 
to  be  accompanied  by  the  liberation  of  certain  substances,  the  activity  of 
which  results  in  profound  changes  in  the  somatic  tissues.  It  is  not  unrea- 
sonable to  suppose  that  the  effect  of  these  substances  upon  the  ovary  might 
exert  equally  important  changes  in  the  egg  apparatus  or  developing  ovules. 
In  its  present  state  this  investigation  appears  to  be  of  great  potential  impor- 


88     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OF   THE   OENOTHERAS. 

tancc,  since  it  offers  the  suggestion  that  we  may  become  able,  more  or  less  at 
will,  to  modify  existing  streams  of  heredity  and  cause  new  organisms  to  arise. 
The  point  not  to  be  lost  sight  of  is  that  the  action  of  external  agents  as 
described  and  deemed  possible  is  not  through  the  somatic  tissues  to  the  egg  or 
germ-plasm,  but  is  exerted  directly  upon  the  reproductive  elements  themselves. 
To  assume  that  such  action  is  stimulative,  as  has  been  done  by  Tower  in  dis- 
cussing the  earlier  announcement  of  these  results  (Tower,  1906),  is  to  adhere  to 
one  of  a  number  of  allowable  suppositions — one,  however,  which  seems  most 
applicable  to  the  results  secured  by  him  with  beetles,  but  one  to  which  we  are 
by  no  means  confined  in  the  consideration  of  the  induction  of  the  atypic  forms 
in  plants.  The  determination  of  the  character  of  this  action  constitutes  one  of 
the  most  difficult  problems  in  connection  with  the  entire  matter. 


MUTATIONS,    VARIATIONS,    AND   RE;i.ATIONSHIPS   OP   THE    OENOTHERAS.      89 

RECAPITULATION. 

The  principal  features  of  the  foregoing  paper  may  be  stated  in  the  following 
brief  generalizations : 

(i)  The  probabihty  of  the  origination  of  new  elementary  strains  of  plants 
arising  by  saltations  or  sporting  is  one  which  has  received  the  support  of  many 
authors,  from  inferential  conclusions  based  upon  the  examination  of  living 
material.  It  has  also  been  found  by  more  than  one  worker  that  characters 
acquired  in  a  saltatory  manner  are  not  swamped  by  intercrossings.  If  such 
characters  behave  as  units  they  may  well  survive,  whether  dominant  or  reces- 
sive with  respect  to  the  corresponding  character  of  the  parental  stock,  the  prob- 
abilities for  survival  being  somewhat  greater  with  recessive  characters. 

(2)  Frequency  of  mutation  is  put  forward  as  a  better  expression  than 
periodicity  of  mutation  for  the  proportion  of  salts  occurring  in  the  course  of 
generations. 

(3)  An  evening- primrose  inseparable  from  0.  lamarckiana  occurred  around 
Haarlem,  Holland,  as  early  as  1756,  and  near  Liverpool,  England,  in  1806. 
It  has  been  noted  in  this  and  other  districts  at  various  times,  and  has  been 
under  more  or  less  continuous  observation  since  1892.  Apparently  estab- 
lished and  growing  with  it  are  0.  rubrinervis  and  0.  lata,  two  of  its  mutants. 
0.  hrevistylis,  another  mutant  which  is  recessive  when  crossed  with  O.  la- 
marckiana, has  been  in  existence  in  competition  with  the  parental  form  in 
Holland  for  twenty  years. 

(4)  The  coefficient  of  mutability  of  Oenothera  lamarckiana  has  not  been 
increased  in  the  cultures  in  America,  although  a  form  which  has  hitherto 
escaped  observation  was  secured.  Seeds  of  this  parental  form,  from  various 
sources  outside  of  Amsterdam,  furnished  a  smaller  proportion  of  mutants  than 
the  material  furnished  by  Professor  De  Vries. 

(5)  0.  ohlonga  has  been  found  to  constitute  as  much  as  54  per  cent  of  the 
atypic  derivatives  of  0.  lamarckiana.  0.  lata,  from  material  grown  in  England, 
was  found  capable  of  self-fertilization  and  gave  rise  to  a  progeny  containing 
the  same  elements  as  when  fertilized  by  the  parental  form. 

(6)  Fixed  hybrids  constituting  species  were  secured  in  combinations  of 
0.  lamarckiana  and  0.  cruciata. 

(7)  The  greater  variability  of  phylogenetically  new  characters  as  compared 
with  older  ones,  which  was  supported  by  statistical  evidence  brought  forward 
in  a  previous  paper,  is  confirmed  by  the  evidence  presented  by  further  studies. 

(8)  The  forms  which  would  appear  to  promise  most  of  importance  in  a 
pedigree-culture  would  be  species  with  a  low  degree  of  variability,  having 
nearly  related  forms  with  a  high  degree  of  variability. 

(9)  The  hypanthium  of  Oenothera,  which  seems  to  be  a  recent  development, 
is  more  variable  than  any  other  feature  of  the  bud. 


90     MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   OI?  THEj   OEJNOTHKRAS. 

(lo)  The  inference  seems  justifiable  that  individual  species  become  less 
variable  as  they  grow  older,  and  that  this  decrease  is  compensated  for  by  the 
fact  that  when  a  new  species  springs  into  existence,  it  has  a  greater  degree  of 
fluctuability  than  its  parent.  Such  saltations  are  the  starting-points  from 
which  series  with  decreasing  fluctuation  follow. 

(ii)  0.  parviflora,  which  has  been  known  in  Europe  since  1759,  has  been 
found  in  its  native  habitat  in  America  in  the  same  manner  that  O.  grandiflora 
was  traced  to  its  original  habitat.  This  leads  to  the  hope  that  O.  lamarckiana 
may  be  found  in  a  wild  state. 

(12)  The  species  of  evening-primrose  of  eastern  America  are  distributed 
in  such  manner  that  the  most  closely  related  species  overlap  in  distribution. 
This  conclusion  is  true,  no  matter  upon  what  taxonomic  basis  the  forms 
are  classified.     Similar  conditions  in  Crataegus  and  Opuntia  are  cited. 

(13)  Two  bud-sports  have  been  followed  through  two  generations  and 
found  to  be  constant.  The  sport  in  one  case  embodied  the  recessive  char- 
acters of  a  hybrid  combination  showing  alternative  inheritance,  the  recessive 
qualities  being  thus  activated  and  extracted  in  the  first  generation,  coming 
true  thereafter. 

(14)  The  action  of  reagents  having  an  osmotic  and  a  chemical  effect  has 
resulted  in  the  induction  of  mutants  in  the  progeny  of  Raimannia  odorata 
and  Oenothera  biennis.  The  mutants  thus  induced  have  been  tested  to  the 
second  and  third  generation  and  found  to  come  true  to  their  newly  assumed 
characters. 

(15)  The  induction  of  mutants  by  the  action  of  reagents  is  a  conclusive 
demonstration  of  the  fact  that  hereditary  characters  may  be  altered  by 
external  forces  acting  directly  upon  the  reproductive  mechanism.  The 
action  of  the  reagents  used  experimentally  is  simulated  by  many  conditions 
occurring  in  nature. 


MUTATIONS,    VARIATIONS,    AND  RELATIONSHIPS   Olf  THE   OENOTHERAS.     Qt 


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